Wetlands of British Columbia: a Guide to Identification

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Wetlands and related ecosystems are vital components of British Columbia's ecological diversity.This useful and beautiful guide presents descriptions of more than 100 wetland, floodplain, estuarine, shallow-water, and "transitional" site associations and their plants, wildlife, and soils. It provides a common language to describe wetland ecosystems and also provides an ecological basis for the management of wetlands. Colour photographs illustrate each of the associations in the fact-sheet summary that outlines essential environmental and biological attributes.

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Please enclose this form along with your certified/business cheque or money order made payable to the “Minister of Finance”. Check out our website at www.publications.gov.bc.ca LAND 52 MANAGEMENT HANDBOOK Wetlands Wetlands of British Columbia

A GUIDE TO IDENTIFICATION of British Columbia A GUIDE TO IDENTIFICATION

Ministry of Forests Forest Science Program Wetlands of British Columbia A GUIDE TO IDENTIFICATION

William H. MacKenzie Jennifer R. Moran

Ministry of Forests Forest Science Program The use of trade, ﬁrm, or corporation names in this publication is for the information and convenience of the reader. Such use does not constitute an ofﬁcial endorsement or approval by the Government of British Colum- bia of any product or service to the exclusion of any others that may also be suitable. Contents of this report are presented as information only. Funding assistance does not imply endorsement of any statements or information contained herein by the Government of British Columbia.

National Library of Canada Cataloguing in Publication Data MacKenzie, William H. Wetlands of British Columbia : a guide to identification (Land management handbook ; 52) Includes bibliographical references: p. ISBN 0-7726-5066-7 1. Wetlands -- British Columbia - Identification. 1. Wetlands -- British Columbia - Classification. I. Moran, Jennifer R. II. British Columbia. Forest Science Program. III. Title. IV. Series. QH87.3.M32 2004 333.91'811'09711 C2003-960224-9

Citation MacKenzie, W.H. and J.R. Moran. 2004. Wetlands of British Columbia: a guide to identification. Res. Br., B.C. Min. For., Victoria, B.C. Land Manage. Handb. No. 52. For more information on Forest Science Program publications, visit our website at www.for.gov.bc.ca/hfd/pubs/index.htm © 2004 Province of British Columbia When using information from this or any Forest Science Program report, please cite fully and correctly.

ii ACKNOWLEDGEMENTS

This field guide could not have been completed without the assistance of many. The “silver-backs” of the provincial ecology program (Jim Pojar, Del Meidinger, Dennis Lloyd, Ray Coupé, Allen Banner, Craig Delong) were as solid ground amid the quaking mires in support of this effort. The initial vision of Allen Banner sparked like a beacon on the misty moors of ignorance. Jim Pojar and Del Meidinger were wise men leading through the shin-tangle of classification. Fieldwork was assisted by many individuals including Candice Ford, Janice Hooge, Kristi Iverson, Larrisa Puls, Danielle Colbert, Arman Mirza, and Dennis’ Angels (Dan Bissonnette, Scott Black, Nicole Brand, Mona Doney, Shane Ford, Kim Johnston, Vanessa Larson, Jessica MacDonald, David Miège and Mike Ryan). Invaluable review comments and material were provided by Marty Kranabetter, Jim Pojar, Del Meidinger, Ray Coupé, Kristi Iverson, Carmen Cadrin, Lisa Mahon, and Reg Newman. The layout of this document was initially inspired by Harris et al. (1996) Field Guide to the Wetland Ecosystem Classification for Northwest Ontario. Paul Dwyer and others at Fractal Design assisted in the design of this document. Paul Nystedt and the staff of the Production Resources Sec- tion helped guide this project to completion. The book was designed by Anna Gamble, typeset by Donna Lindenberg of Newport Bay Publishing Limited, and edited and proofread by Steven Justin Smith. Funding was generously provided by Forest Renewal BC, the Ministry of Forests Research Branch, and the Ministry of Environment Habitat Inventory Branch.

iii

CONTENTS

Acknowledgements ...... iv 1 Introduction...... 1 Other Sources of Information ...... 2 Guide Content and Limitations ...... 3 2 Ecology and Classification of Wetland Ecosystems ...... 5 Characteristics of Wetlands and Related Ecosystems ...... 6 A Synopsis of the Classification System...... 11 Classification of Wetland and Riparian Soils ...... 16 Regional Classifications...... 17 Summary of Sampling and Analysis Methods ...... 22 3 Procedures for Site Description and Identification ...... 25 Describing and Identifying Site Units ...... 26 4 Site Classes ...... 29 Key to Site Classes ...... 30 Brief Descriptions of Site Classes ...... 30 5 Site Associations ...... 37 Layout and Conventions for Site Class Descriptions ...... 38 Site Association Fact Sheet ...... 39 Taxonomic Considerations...... 46 5.1 Bogs ...... 49 Wb01 Black spruce – Creeping-snowberry – Peat-moss ...... 56 Wb02 Lodgepole pine – Bog rosemary – Peat-moss ...... 57 Wb03 Black spruce – Lingonberry – Peat-moss...... 58 Wb04 Western hemlock – Cloudberry – Peat-moss ...... 59 Wb05 Black spruce – Water sedge – Peat-moss ...... 60 Wb06 Tamarack – Water sedge – Fen moss ...... 61 Wb07 Lodgepole pine – Water sedge – Peat-moss...... 62 Wb08 Black spruce – Soft-leaved sedge – Peat-moss...... 63 Wb09 Black spruce – Common horsetail – Peat-moss ...... 64 Wb10 Lodgepole pine – Few-flowered sedge – Peat-moss . . . . . 65 Wb11 Black spruce – Buckbean – Peat-moss ...... 66 Wb12 Scheuchzeria – Peat-moss ...... 67 Wb13 Shore sedge – Buckbean – Peat-moss ...... 68 Wb50 Labrador tea – Bog-laurel – Peat-moss ...... 69

v Wb51 Shore pine – Black crowberry – Tough peat-moss ...... 70 Wb52 Common juniper – Tufted clubrush – Hoary rock-moss. . 71 Wb53 Shore pine – Yellow-cedar – Tufted clubrush ...... 72 Additional Bog Site Associations ...... 73 5.2 Fens...... 75 Wf01 Water sedge – Beaked sedge...... 82 Wf02 Scrub birch – Water sedge ...... 83 Wf03 Water sedge – Peat-moss ...... 84 Wf04 Barclay’s willow – Water sedge – Glow moss ...... 85 Wf05 Slender sedge – Common hook-moss ...... 86 Wf06 Slender sedge – Buckbean ...... 87 Wf07 Scrub birch – Buckbean – Shore sedge...... 88 Wf08 Shore sedge – Buckbean – Hook-moss...... 89 Wf09 Few-flowered spike-rush – Hook-moss ...... 90 Wf10 Hudson Bay clubrush – Red hook-moss...... 91 Wf11 Tufted clubrush – Star moss ...... 92 Wf12 Narrow-leaved cotton-grass – Marsh-marigold ...... 93 Wf13 Narrow-leaved cotton-grass – Shore sedge ...... 94 Wf50 Narrow-leaved cotton-grass – Peat-moss ...... 95 Wf51 Sitka sedge – Peat-moss ...... 96 Wf52 Sweet gale – Sitka sedge ...... 97 Wf53 Slender sedge – White beak-rush ...... 98 Additional Fen Site Associations ...... 99 5.3 Marshes ...... 101 Wm01 Beaked sedge – Water sedge...... 108 Wm02 Swamp horsetail – Beaked sedge ...... 109 Wm03 Awned sedge ...... 110 Wm04 Common spike-rush ...... 111 Wm05 Cattail ...... 112 Wm06 Great bulrush ...... 113 Wm07 Baltic rush ...... 114 Wm50 Sitka sedge – Hemlock-parsley ...... 115 Wm51 Three-way sedge ...... 116 Additional Marsh Site Associations...... 117 5.4 Swamps ...... 121 Ws01 Mountain alder – Skunk cabbage – Lady fern ...... 128 Ws02 Mountain alder – Pink spirea – Sitka sedge ...... 129 Ws03 Bebb’s willow – Bluejoint...... 130

vi Ws04 Drummond’s willow – Beaked sedge ...... 131 Ws05 MacCalla’s willow – Beaked sedge...... 132 Ws06 Sitka willow – Sitka sedge ...... 133 Ws07 Spruce – Common horsetail – Leafy moss ...... 134 Ws08 Subalpine fir – Sitka valerian – Common horsetail...... 135 Ws09 Black spruce – Skunk cabbage – Peat-moss ...... 136 Ws10 Western redcedar – Spruce – Skunk cabbage ...... 137 Ws11 Spruce – Subalpine fir – Skunk cabbage ...... 138 Ws50 Pink spirea – Sitka sedge ...... 139 Ws51 Sitka willow – Paciﬁc willow – Skunk cabbage ...... 140 Ws52 Red alder – Skunk cabbage...... 141 Ws53 Western redcedar – Sword fern – Skunk cabbage ...... 142 Ws54 Western redcedar – Western hemlock – Skunk cabbage. . . 143 Ws55 Yellow-cedar – Mountain hemlock – Skunk cabbage . . . . 144 Additional Swamp Site Associations ...... 145 5.5 Shallow-waters...... 147 5.6 Estuarine Associations ...... 155 Em01 Widgeon-grass ...... 162 Em02 Glasswort – Sea-milkwort ...... 163 Em03 Seashore saltgrass ...... 164 Em04 Seaside plantain – Dwarf alkaligrass...... 165 Em05 Lyngbye’s sedge...... 166 Em06 Lyngbye’s sedge – Douglas’ water-hemlock ...... 167 Ed01 Tufted hairgrass – Meadow barley ...... 168 Ed02 Tufted hairgrass – Douglas’ aster ...... 169 Ed03 Arctic rush – Alaska plantain ...... 170 Additional Estuarine Site Associations ...... 171 5.7 Flood Associations ...... 173 Fl01 Mountain alder – Common horsetail ...... 180 Fl02 Mountain alder – Red-osier dogwood – Lady fern ...... 181 Fl03 Paciﬁc willow – Red-osier dogwood – Horsetail ...... 182 Fl04 Sitka willow – Red-osier dogwood – Horsetail ...... 183 Fl05 Drummond’s willow – Bluejoint ...... 184 Fl06 Sandbar willow ...... 185 Fl07 Water birch – Rose ...... 186 Fm01 Cottonwood – Snowberry – Rose ...... 187 Fm02 Cottonwood – Spruce – Red-osier dogwood...... 188 Fm03 Cottonwood – Subalpine fir – Devil’s club ...... 189

vii Fl50 Sitka willow – False lily-of-the-valley ...... 190 Fl51 Red alder – Salmonberry – Horsetail ...... 191 Fm50 Cottonwood – Red alder – Salmonberry ...... 192 Additional Flood Site Associations ...... 193 5.8 “Transition” Associations ...... 195 Gs01 Alkali saltgrass ...... 202 Gs02 Nuttall’s alkaligrass – Foxtail barley...... 203 Gs03 Field sedge ...... 204 Gs04 Tufted hairgrass ...... 205 Sc01 Scrub birch – Kinnikinnick ...... 206 Sc02 Grey-leaved willow – Glow moss ...... 207 Sc03 Barclay’s willow – Arrow-leaved groundsel ...... 208 6 Conservation and Management Issues ...... 209 Livestock Management ...... 210 Timber Harvest ...... 213 Wetlands as Wildlife Habitat ...... 217  1 Ordination of Site Associations ...... 239 2 Overlays of the Wetland Edatopic Grid...... 242 3 Wetland Indicator Plant Species ...... 245 4 Crosswalk to Other Classiﬁcations...... 249 5 Scientific names with common names ...... 254 6 Wildlife Species Afﬁliated with Wetlands and Related Ecosystems...... 261 7 Glossary...... 269 Literature Cited ...... 279  3.1. Important site and soil features for identifying site units...... 27 4.1. Summary of characteristics for wetland and related ecosystem Site Classes ...... 31 5.1.1 Distribution of Bog Site Associations by biogeoclimatic zone. . . 53 5.1.2 Bog Species Importance Table ...... 54 5.2.1 Distribution of Fen Site Associations by biogeoclimatic zone . . . 79 5.2.2 Fen Species Importance Table...... 80 5.3.1 Distribution of Marsh Site Associations by biogeoclimatic zone ...... 105 5.3.2 Marsh Species Importance Table ...... 106

viii 5.4.1 Distribution of Swamp Site Associations by biogeoclimatic zone ...... 125 5.4.2 Swamp Species Importance Table...... 126 5.5.1 Characteristics of water with different nutrient status ...... 149 5.6.1 Distribution of Estuarine Site Associations by biogeoclimatic zone...... 159 5.6.2 Estuarine Species Importance Table...... 160 5.7.1 Distribution of Flood Site Associations by biogeoclimatic zone ...... 177 5.7.2 Flood Species Importance Table ...... 178 5.8.1 Distribution of “Transition” Site Associations by biogeoclimatic zone...... 199 5.8.2 “Transition” Species Importance Table ...... 200 6.1 Distribution and habitat use by adult pond-breeding amphibians ...... 219 6.2 Distribution and habitat use by dabbling ducks ...... 226 6.3 Distribution and habitat use by “lake” ducks ...... 228 6.4 Distribution and habitat use by cavity-nesting ducks...... 229 A3.1 List of Obligate Hydrophytes...... 245 A3.2 List of Facultative Hydrophytes – Wetland Affiliated ...... 246 A3.3 List of Facultative Hydrophytes – Upland Affiliated...... 248 A4.1 Wetland and related ecosystem Site Associations with corresponding BEC site series...... 249 A4.2 Wetland Site Associations corresponding to the Cariboo wetland classification...... 250 A4.3 Wetland Site Associations corresponding with Klinka et al. 1997 ...... 251 A4.4 Wetland Site Associations corresponding with coastal classifications in Washington and Alaska...... 252 A4.5 Wetland Site Associations corresponding with classifications in Washington and Montana ...... 253  2.1 Site unit hierarchy ...... 13 2.2 Conceptual relationship between ecosystem Realms ...... 15 2.3 Examples of coding for Site Associations...... 15 2.4 Biogeoclimatic zones of British Columbia ...... 18 2.5 Ecoprovinces of British Columbia ...... 20 4.1 Dichotomous key for the identification of Site Class ...... 32 4.2 Site Class distribution on the modified edatopic grid...... 33 5.0.1 Example Site Association fact sheet ...... 40

ix 5.0.2 A modified edatopic grid showing pH and hydrodynamic index axes for Wet and Very Wet sites ...... 42 5.0.3 Environmental characteristics useful for determining nutrient status in wetlands...... 43 5.0.4 Distribution of Site Classes relative to intertidal zone and salinity ...... 45 5.1.1 Position of bogs on the edatopic grid ...... 51 5.2.1 Position of fens on the edatopic grid ...... 77 5.3.1 Position of marshes on the edatopic grid...... 103 5.4.1 Position of swamps on the edatopic grid ...... 123 5.6.1 Distribution of ecosystem classes of estuaries by elevation and salinity...... 157 5.7.1 Position of flood ecosystems on the edatopic grid ...... 175 5.8.1 Position of “transition” ecosystems on the edatopic grid ...... 197 A1.1  ordination of numbered wetland and related Site Associations including some shallow-water units ...... 240 A1.2  ordination of numbered wetland Site Associations only. . . 241 A2.1 Edatopic grid position of vegetation physiognomy...... 242 A2.2 Edatopic grid position of major bryophyte groups and peat-accumulating sites ...... 243 A2.3 Edatopic grid position of Classes from the British Columbia Vegetation Classification ...... 244

x Introduction 1

Wetlands are relatively uncommon ecosystems throughout most of British Columbia and yet play an integral role. Montane ponds and fens, such as this one near Alice Arm, support high- elevation wildlife populations and regulate headwater hydrology. This guide presents a site classification and interpretative information for wetlands and related ecosystems of British Columbia. Site identification is based upon principles of Biogeoclimatic Ecosystem Classification () modified for wetland ecosystems. The objectives of the classification are: • to provide a framework for organizing ecological information and management experience about ecosystems; • to promote a better understanding of wetlands and related ecosystems; • to provide users with a common language to describe wetland ecosystems; and • to provide an ecological basis for management of wetlands. The Research Branch of British Columbia’s Ministry of Forests initiated a program in 1994 to classify and describe the wetlands and riparian areas of British Columbia. Its central intent was to generate basic ecological information about these important ecosystems. This guide represents one aspect of the Wetland and Riparian Ecosystem Classifica- tion () initiative and has two principal goals: • to assist users in describing and identifying wetland ecosystems; and • to provide management interpretations to assist in conservation of wetlands ecosystems.

OTHER SOURCES OF INFORMATION

A more detailed description of the wetland classification framework can be found in Classification of Wetlands and Related Ecosystems in British Columbia (MacKenzie and Banner 2001). Explanations of the national wetland classification are presented in the Wetlands of Canada ( 1988). Biogeoclimatic Ecosystem Classification in British Columbia (Pojar et al. 1987) outlines the methods and philosophy of ; descriptions of biogeoclimatic zones and representative ecosystems can be found in Ecosystems of British Columbia (Meidinger and Pojar 1991). For more detailed discussion of the standard methods used to describe ecosystems in British Columbia, refer to Describing Ecosystems in the Field (2nd edi- tion) (Luttmerding et al. 1990), Field Manual for Describing Terrestrial Ecosystems (Province of British Columbia 1998), and A Wetland Sampling Methodology: Draft (MacKenzie and Shaw 1998). Regional treatments of wetland Site Associations have been published for the Cariboo (Steen and Roberts 1988), south Coast (Klinka et al.

2 Wetlands of British Columbia: A Field Guide to Identification unpublished), and north Coast (Banner et al. 1986). Regional  field- guides (e.g., Banner et al. 1993) also contain descriptions for some forested swamp, bog, and flood ecosystems. For a comparison to wetland units described in other classifications, refer to Appendix 4. For field identification of wetland plant species, refer to any of the Lone Pine series. For example, Plants of Northern British Columbia (MacKin- non et al. 1992) can be helpful for willow identification and Plants of the Western Boreal Forest and Aspen Parkland (Johnson et al. 1995) is good for sedges and mosses, especially Sphagnum species. The Illustrated Flora of British Columbia (Douglas et al. 1998–2002) provides formal botanical keys to vascular species identification.

GUIDE CONTENT AND LIMITATIONS

This guide describes common wetland ecosystems that occur in British Columbia. Related types of ecosystems that do not meet the strict definition for true wetlands are also presented. These include: • Flood ecosystems, which occur on the floodplains of rivers and are inundated during the spring freshet but have well-drained and aerated soils; •“Transition” ecosystems, which are often associated with wetlands and can have the vegetation structure similar to a wetland but have merely moist soils and few wetland indicator species; and • tidal Estuarine ecosystems, which occur at the confluence of marine and fluvial environments. Alpine wetlands are not described in this guide. No guide can encompass all of the complexity and diversity of ecosystems that occur on the landscape. Although the described units include most of the common wetland and related ecosystems found in the province, users are likely to encounter sites that do not appear to “fit” the classification. Furthermore, wetlands are complex ecosystems that can change rapidly with hydrological modifications. This guide describes ecosystems that recur throughout the landscape and appear to be relatively stable in vegetation composition. An understanding of basic ecological factors and wetland processes is required to assess whether an equivocal site is in transition, represents a distinct ecological community not yet described, or is a variation of a described site association. This guide is just one tool to help describe and compare wetland and related

Chapter 1 Introduction 3 ecosystems and must be supplemented with practical knowledge, experience, and judgement. The guide consists of the following major sections: Chapter 2 provides an overview of the basic ecological features and environments of wetlands and related ecosystems and a summary of the classification systems underlying units in this guide. Chapter 3 explains the taxonomic conventions, format, and coding used throughout the guide and the process of site identification. Chapter 4 gives several tools for identifying the Site Class (broad func- tional groups of wetland ecosystems) including keys, tables of characteristics, and brief descriptive summaries. Chapter 5 constitutes the main body of the guide. This chapter provides basic ecological information for the recognition of different wetland ecosystem types that occur in British Columbia. In standardized fact sheet format, information is provided for all wetland Site Classes and Site Associations. The chapter is divided into eight sections, one for each Site Class or Group described in this guide. A description of the major ecological characteristics for each recognized Association is presented in a one-page fact-sheet format outlining site, vegetation, and environmental characteristics. Short descriptions are included at the end of each section for infrequently sampled or poorly known ecosystems. Chapter 6 summarizes conservation and management issues for wetlands and related ecosystems. This information will allow resource managers to make better-informed decisions on conservation and management of these ecosystems. The appendices provide background information for the guide, including details about the multivariate analysis used to create the classification, a crosswalk to similar previously described units, a list of hydrophytes, a list of wetland-affiliated wildlife species, a glossary, and interpretive overlays of the wetland edatopic grid.

4 Wetlands of British Columbia: A Field Guide to Identiﬁcation Ecology and 2 classiﬁcation of wetland ecosystems

The habitats surrounding wetlands are essential for a functioning wetland ecosystem. Most wildlife inhabiting this marsh near Lac La Hache will also use the adjacent grassland, aspen copse, or coniferous forest for some important part of their life cycle. This section brieﬂy describes the important ecological themes and principal concepts in ecosystem classiﬁcation of wetland and related ecosystems.

CHARACTERISTICS OF WETLANDS AND RELATED ECOSYSTEMS

Wetland Ecosystems

Wetlands are: areas where soils are water-saturated for a sufficient length of time such that excess water and resulting low soil oxygen levels are principal determinants of vegetation and soil development. Wetlands will have a relative abundance of hydrophytes in the vegetation community and/or soils featuring “hydric” characters. This wetland definition encompasses a wide range of ecosystems, from semi-terrestrial fens, bogs, and swamps to semi-aquatic marshes and shallow open water. Wetlands include a broad range of ecosystem types, from those permanently flooded by shallow water and dominated by aquatic organisms to forested sites with merely wet soils. The water-saturated environment of wetlands supports a unique group of plants called hydrophytes. These plants are adapted to grow in waterlogged soils. Excessive water and the low rate at which oxygen diffuses under these conditions leads to a complex of critical conditions that require specialized adaptations (Daubenmire 1959). Adaptations, such as leathery leaves (to reduce nutrient requirements and combat physiologi- cal drought) or specialized internal air compartments (to transport oxygen to the roots) are required for wetland plants. Obligate hydrophytes (such as great bulrush) are restricted to wetlands and semi-aquatic sites. Facultative hydrophytes (such as Labrador tea and many other members of the Heather family) occur commonly in wetlands but also appear on some upland sites. Wetland soils are subhydric or hydric and have one or more of the following features that reflect anaerobic soil conditions: 1. Peaty organic horizons greater than 40 cm thick. 2. Non-sandy soils with blue-grey gleying within 30 cm of the surface. 3. Sandy soils with prominent mottles within 30 cm of the surface or blue-grey matrix. 4. Hydrogen sulphide (rotten egg smell) in upper 30 cm.

6 Wetlands of British Columbia: A Field Guide to Identification From an ecological perspective, either an abundance of hydrophytes or hydric soil conditions is generally sufficient to indicate a wetland ecosystem. The boundary of the wetland is identified by changes in vegetation structure, loss of hydrophytes, and absence of wetland soil characteristics. Environmental gradients in wetland ecosystems The stability, mobility, and chemical composition of the watertable are the major environmental variables at the site level that differentiate wetland ecosystems. The acidity/alkalinity of the watertable relates directly to base cation content and indirectly to nutrient availability in wetland ecosystems. Major floristic shifts occur over this gradient, particularly in peatlands (Vitt 1994). Ombrotrophic sites receive surface water from precipitation only and have few available base cations, high acidity, and very poor nutrient regime, and are dominated by Sphagnum mosses. Where the surface waters have been in contact with mineral materials, base cations in the surface water increase and the site becomes more alkaline, and peatlands become dominated by brown mosses (e.g., Campylium, Tomenthypnum, Calliergon, Drepanocladus species). Alkali are mineral soils that contain an excess of exchangeable sodium and pH >8.5. Alkali soils will often have impermeable clay horizons and dry to a dark crust. Saline soils do not occur in peatlands but in saline meadows where evaporation concentrates salts. Saline sites have excess soluable salts (Na+, Ca++, Mg++, Cl-, SO4--) that impair site productivity, pH < 8.5, and dry to a white crust. Saline-alkali soils also occur. These sites have impermeable clay horizons, pH >8.5, and dry to a white crust (Richards 1969). Regional bedrock and surface geology, and site drainage and discharge, influence the base cation content of the groundwater supply. For example, groundwater percolation in the Rocky Mountains generally has very high content of base cation (calcium) because of contact with limestone parent materials. Rich fens and swamps are common in this area. The outer Coast is dominated by nutrient-poor bogs largely due to the sterile nature of the granitic bedrock that underlies this region. Water movement and seasonal water-level fluctuation (hydrodynamics) are as important as soil moisture regime in wetland ecosystems. Soil moisture regime can be difficult to assess in many wetland and flood

Chapter 2 Ecology and classification of wetland ecosystems 7 sites because the watertable changes significantly over the growing season. As a general rule, sites with greater waterflow are richer. Increasing lateral movement of water improves nutrient availability by bringing additional supplies of minerals and improving oxygenation. Vertical movements of the watertable alternately flood and expose the surface of the wetland, improving aeration and increasing decomposition rates. Stable, high watertables (stagnant or sluggish hydrodynamics) promote peat formation and high bryophyte cover. Bogs and fens are ecosystems that form under these conditions. Peat accumulations on these sites can be very deep, are usually poorly decomposed, and are largely derived from mosses and sedges. Sites with more dynamic watertables experience surface flooding followed by late-season drawdown. These types of sites usually have few bryophytes because most mosses are intolerant of prolonged submer- gence. Marshes, swamps, and shallow-water ecosystems may be underlain with peat but it is usually well-humified, non-bryophytic material derived from sedges or wood. On these sites and on flood ecosystems, the length and depth of flooding and the degree of waterflow are primary factors determining community composition. Species composition depends on species flood tolerance and life history requirements. Sites with large fluctuations in watertable, such as pothole marshes or tidal marshes, represent some of the most hydrologically dynamic and nutrient-rich ecosystems. Regional climate is the broad environmental context for ecosystems. For wetlands, this influence is less pronounced than for upland ecosystems because of the overriding influence of wet site conditions. Still, climate is an important factor in wetland formation. In cool climates, where water inputs exceed evaporation rates, peatland formation is promoted. Small, water-collecting depressions and lake margins that would be marshes in regions with warm, dry climates are topogenous peatlands here. In these peatland regions, only the most hydrologically active areas adjacent to rivers and large lakes support marshes and shrub swamps. Peatlands occur even on sloping terrain in regions of cool climates with high summer precipitation, extensive groundwater seeps, or prolonged snowpack, where continual groundwater inputs maintain surface saturation. The dry and warm regions of the province typically have very limited peatland development because warm temperatures increase decomposition rates and high evaporation rates limit the extent of sites that have permanent saturation.

8 Wetlands of British Columbia: A Field Guide to Identification Estuarine ecosystems Estuarine ecosystems are defined as: coastal sites dominated by plants and other organisms tolerant of wet, brackish soils, found at the confluence of a freshwater source and the marine environment and affected by occasional or diurnal tidal inundation. Estuarine ecosystems occur specifically where: • at least periodically, the land supports predominantly hydrophytic plant species or benthic fauna adapted to brackish water. • the substrate is predominantly undrained hydric soil. Soils may be organic or inorganic. In mineral soils, gleying occurs within the top 30 cm, or the substrate is nonsoil and is saturated with water or covered by shallow water at some time during the growing season. • the site is tidally influenced and at least occasionally affected by brackish water. Estuarine ecosystems have similar characteristics to wetland ecosystems, with the additional influences of diurnal fluctuations in watertable and variable salinity. The gradients of most importance in this realm of ecosystems is the degree of tidal flooding, which is closely related to height above the mean tide level. Ecosystems that occur at the lowest level are flooded with every tide (excepting neap tides), while the highest may experience only occasional flooding during the highest high tides. The degree of freshwater influence affects species distribution within an estuary independent of other factors. Particularly where high volumes of fresh water are delivered to estuarine environments, communities change along a gradient from where freshwater influences predominate (usually within the tidal reaches of the river) to where freshwater inputs are minimal. Low-elevation coastal climates are more equable than interior and high- elevation climates, yet climate still plays a role in estuarine development. Estuaries of the Georgia Depression, where summers are warm and dry and where freshwater inflows are highest in the winter and lowest in the summer, have a flora that is related to estuaries farther south. Estuaries of the Coast and Mountains, where summers are cool and moist and rivers have peak flows in spring and summer, lack many of the “Cali- fornian” estuary species. Terrestrial ecosystems Terrestrial ecosystems occur on sites where water is not in surplus for extended periods of the growing season (actual moisture regime Very Dry

Chapter 2 Ecology and classification of wetland ecosystems 9 to Very Moist). These sites are characterized by the dominance of drought-tolerant vascular plants, bryophytes, and cryptogams. Decom- position is primarily aerobic by fungi, bacteria, and soil fauna. Within a climatic region, differentiation of most terrestrial ecosystems can be explained primarily by soil moisture regime () and soil nutrient regime (). However, the two groups of terrestrial ecosystems included in this guide, flood and “transition,” have special environmental factors that override  and  influences. The term “riparian” is defined or applied in the literature in several con- tradictory ways. In , this term is used to describe sites that are adjacent to a waterbody. Thus, “riparian” does not imply any specific ecological feature of a site other than occurence next to a waterbody. By this definition, any type of ecosystem can be riparian. For those ecosystems that are ecologically distinct because of flooding, erosion/ sedimentation, or subirrigation from an adjacent waterbody, this guide uses the term “Flood ecosystem.” Flood ecosystems are defined as: sites flooded for a short time during the growing season, where soils are freely drained and anoxic conditions (if they occur) are quickly relieved after subsidence of floodwater. Vegetation tolerant of brief flooding events but not prolonged soil saturation is typical. Flood ecosystems occur specifically where: • waterbodies periodically flood their banks, depositing or eroding fluvial or lacustrine materials; and • watertables are within the rooting zone during part of the growing season, but not for sufficient duration to cause gleying within the top 30 cm of soil depth. The duration and power of flooding are the primary site determinants in these ecosystems. “Transition” ecosystems are a loosely defined set of Associations that are included in this guide because they often occur adjacent to wetlands and have some structural and ecological similarities. Two Classes are described, the saline meadows of the Grassland Group and shrub-carrs from a Shrubland Group. Both of these Classes have Moist or Very Moist actual moisture regime and support non-forested climax communities. Because of a special site factor, these sites do not support forest or grassland ecosystem species groups that would normally occur under these soil moisture regimes.

10 Wetlands of British Columbia: A Field Guide to Identification Saline meadows have high soil salinity or alkalinity. They occur only in dry climates where early-season flooding prevents growth of trees or upland grasses, but where sufficient drawdown occurs to limit hydrophyte establishment. High content of salts in the soil is the driving variable for these meadows. Shrub-carrs only occur in cold climates where cold-air drainage or ponding on some sites results in cold soils and mid-season frosts that preclude tree establishment. These areas are primarily in the Interior at high elevations or in boreal climates.

A SYNOPSIS OF THE CLASSIFICATION SYSTEM

Concepts This guide applies a site unit classification model of a Wetland and Ri- parian Ecosystem Classification system () (MacKenzie and Banner 2001). This system has its basis in the Biogeoclimatic Ecosystem Classifi- cation () (Pojar et al. 1987) and Canadian Wetland Classification System () (Warner and Rubec 1997). Ecosystem Ecosystems are interacting complexes of living organisms and their physical/chemical environments. For purposes of this guide, ecosystems are defined as portions of the physical landscape and the living systems that are on and in it. They are identified and characterized by a plant community and its associated environment. More specifically, ecosystems are areas of relatively uniform vegetation, topography, soils, and hydrology. Microtopographic features are an important environmental factor in wetland ecosystems. Several distinct plant communities can occur on the hummocks and in the hollows of a single site. In this guide, these fine- scale variations are not treated as separate units but are considered to be a normal (and predictable) state for the ecosystem. Climax, succession, and site potential  integrates vegetation, climatic, site, and seral classifications into a system of regional, local, and chronological units. The basic approach in  is to use mature or climax vegetation communities to define site and climate classification units. “Climax” in ecology refers to a condition of dynamic equilibrium, a steady state rather than a static endpoint. For vegetation, this means that the species in the community replace them- selves rather than being replaced by other species over time (the process of succession).

Chapter 2 Ecology and classification of wetland ecosystems 11 Mature communities are said to reflect site potential (or climatic potential in the restricted case of a zonal ecosystem) because they are relatively stable and are thought to integrate and reflect the sum of all the environmental conditions of the site. Site potential is a central concept in  but has been applied primarily to forested ecosystems, where the mature state is recognized through canopy structure and species composition. Applying this approach for wetlands and flood ecosystems is more difficult because they represent more dynamic systems where the site potential changes over time. Deviations from the concept of site potential occur in flood ecosystems, where deposition and erosion during flooding change soil texture, depth to watertable, and duration of flooding over time. These changes to site potential may occur gradually over decades or abruptly with the impacts of a significant flood. Similarly, the gradual accumulation of organic matter and establishment of Sphagnum mosses both affect the hydrological properties of peatlands, leading to long-term peatland succession and changes in site potential. Pothole marshes are characteristically variable in their interannual water regime; sites may be flooded in one year and dry in the next. Therefore, the concepts of climax and site potential in wetland and floodplain sites must be viewed as more fluid and temporary than in upland ecosystems. Classification versus description Site description and ecosystem classification are different but complementary processes. Site description produces a simple list of biotic and abiotic features for an ecosystem. No two ecosystems will have the exact same list of site characteristics and each site could be considered unique. However, to apply knowledge gained on one site more widely, groups of sites with similar ecological function must be recognized. Ecosystem classification distills the commonality among sites into recognizable groups based on a few ecologically important factors. These fundamental properties feature prominently within the formal classification; other descriptive attributes are used as supporting information. Site units versus landscape units and mapping The classification presented in this guide is a site classification; it describes the ecosystem as defined in the previous section—an area of relatively uniform vegetation, topography, soils, and hydrology. Howev- er, most wetlands and riparian areas are complexes of ecosystems, where mapping of site units would be difficult at commonly used scales (e.g., 1:20 000). For this reason, it is more likely that entire wetlands will be mapped as a single unit.  proposes a landscape-level unit called the

12 Wetlands of British Columbia: A Field Guide to Identification Ecocomplex to describe recurring complexes of Site Associations for use in mapping wetlands (MacKenzie and Banner 2001). Classification units  integrates several different classification models into a single hier- archical framework. The Site Association unit of  is grouped into broader units on the basis of more general ecological similarities (Figure 2.1). The Site Series and Site Association units of  describe site potential on ecologically homogeneous areas based on climate, soils, and mature vegetation communities. The “Class” concept of the Canadian Wetland Classification System is used as a broader description of site potential. Additional, even broader units (Group and Realm) formalize currently used, but imprecisely defined, terminology. These higher levels are defined by environmental states characterized by specific guilds of biota, rather than species groups. The units accentuate similarities in basic underlying processes and functions between ecosystems not reflect- ed in vegetation. Such broad units allow wetlands to be placed in context with other non-wetland ecosystems. Site Association The Site Association defines all sites capable of supporting a similar plant association at climax. Vegetation classification using a Braun- B Blaunquet approach produces units with characteristic diagnostic species groups (Mueller-Dombois and Ellenberg 1974). E When these species groups represent climax communities, they are said to reflect Series site potential and are used to define the C Site Series within a given biogeoclimatic Association subzone/variant, or a Site Association across several  subzones/variants Class (Pojar et al. 1987). The Site Association is a collection of Site Series with similar plant Group species composition. For example, the Wb01 (Black spruce – Lingonberry – Peat- moss) Site Association includes the Site Realm Series BWBSdk1/10, BWBSdk2/07, BWBSmw1/08, and BWBSmw2/08.  . Site unit hierarchy (from MacKenzie and Banner 2001).

Chapter 2 Ecology and classification of wetland ecosystems 13 Site Class The Site Class describes Associations with similar basic underlying environmental attributes that support similar characteristic species guilds at climax. The Class concept is adapted from the Wetland Class of the Canadian Wetland Classification System () (Warner and Rubec 1997). The Site Class, as applied here, is narrower than the original  concept, where the Class could describe entire wetlands and with no requirement for “climax” plant communities. In this guide, the Class is an extension of the Site Association concept and the same restrictions apply: sites must be relatively homogeneous (rather than entire wetlands), mature, and undisturbed. Site Group 1 The Group describes functionally similar Classes based on a dominant, ecologically relevant environmental feature(s). A single dominant environmental factor or site attribute, reflecting a constellation of environmental factors that influence ecosystem structure, is used to differentiate Groups. For example, within the Wetland Realm, the Peatland Group is distinguished from the Mineral Group based on the presence of deep fibric or mesic peat accumulations that indicate low decomposition rates, lower available nutrients, and near- permanent saturation. The Flood Group of the Terrestrial Realm includes those ecosystems that are strongly influenced by periodic/seasonal flooding events common in riparian situations. Site Realm The Realm delineates major biotic types that reflect gross differences in water abundance, quality, and source. There are three primary Realms (Terrestrial, Freshwater, and Marine) and four secondary Realms where the primary Realms intersect (Wet- land, Estuarine, Intertidal, and Wedge). The secondary Realms exhibit unique characteristics in addition to features that are common to the related primary Realms. This guide covers ecosystems of the wetland and estuarine ecosystem Realms and some terrestrial ecosystem groups that are related to true wetlands (Figure 2.2).

1 The Site Group term was previously used in  to define broad groupings of Site Associations based on species similarity. This  unit is now referred to as the Site Alliance to reflect its basis in vegetation classification.

14 Wetlands of British Columbia: A Field Guide to Identiﬁcation Wetland Terrestrial Freshwater Estuarine Intertidal Wedge

Marine

 . Conceptual relationship between ecosystem Realms. Shaded section shows ecosystems covered in this guide (from MacKenzie and Banner 2001). The shaded portion of the Terrestrial Realm includes flood and “transition” ecosystems.

Naming and numbering of Site Units Classes and Associations are given a two-letter Class and two-number Association code (Figure 2.3). The Class code is a combination of the ﬁrst letter of the Realm or Group in capital plus the ﬁrst letter of the Class in small capitals. The number for the Site Association is always two digits. Interior Associations or those that occur on the Coast but are primarily of Interior distribution are numbered 01 to 49. Site Associations that are exclusively or primarily Coastal are numbered 50 to 99. Num- bering of units loosely follows an approach of assigning the “01” number to an Association considered to represent the central concept for the Class. Units that most closely resemble this initial Association are then numbered sequentially. Note that these Site Association codes are for application in a broader context, where the user intends to discuss ecosystems at a regional or provincial level. For discussion in a local context, the  Site Series code should be used if applicable (e.g., SBSdk/10). Refer to Appendix 4.

W b 01 (Black spruce – Creeping-snowberry – Peat-moss)

Realm: Wetland Class: Bog Site Association #1 (Interior) Group: Flood Class: Low Bench Site Association #1 (Coastal)

F l 50 (Sitka willow – False lily-of-the-valley)

 . Examples of coding for Site Associations.

Chapter 2 Ecology and classiﬁcation of wetland ecosystems 15 Full names of the Association follow the Site Series standards for : no more than three species are used, species from the dominant layers appear ﬁrst.

CLASSIFICATION OF WETLAND AND RIPARIAN SOILS

Soil types used in this guide are from the Canadian System of Soil Classi- fication () (Agriculture Canada Expert Committee on Soil Survey 1987). There are nine Soil Orders in the , of which two are found primarily in wetland environments: Gleysol Order and Organic Order. Gleysols are defined by the presence of a gleyed soil horizon. Gleying occurs under waterlogged, anaerobic conditions. A gleyed horizon is indicated by a distinct dull blue-grey colour or prominent rust-coloured blotches called mottles. Humic Gleysols have an organically enriched surface horizon in addition to the above features. Rego Gleysols are very young and have limited profile development. The soils of the Organic Order include Fibrisols, Mesisols, Humisols, and Folisols (upland organic soils); the first three are wetland soil Great Groups. These soils have accumulated more than 60 cm of organic materials derived from hydrophytic plants. Most organic soils are saturated with water for prolonged periods. They occur widely in poorly and very poorly drained depressions and level areas in regions of cool or wet climates. Classification at the Great Group level (Fibrisol, Mesisol, or Humisol) is based primarily on the dominant horizon of the decomposition of the organic material in the middle tier (between 60 and 160 cm below the soil surface). Fibrisols are poorly decomposed, and peat constituents are easily recognizable. Mesisols are partially decomposed and Humisols are well decomposed. Typic organic soils have >160 cm or organic accumulations while Terric organic soils have 60–160 cm of organic matter over mineral substrates. Several other Orders are described in this guide for non-wetland Site Associations. Regosols are very young soils with little or no horizon development. They are common on active fluvial sites where flood events deposit sediment layers. On these sites, Regosols are cumulic with layers of sediments from different flood events. Brunisols are slightly older soils, with some chemical weathering, and can be gleyed from short periods of saturation.

16 Wetlands of British Columbia: A Field Guide to Identiﬁcation Solonetzic soils have a prismatic soil structure resulting for the accumulation of salts. Saline meadows are classic locations for Solonetzic soils.

REGIONAL CLASSIFICATIONS

Two complementary regional classifications are used in Site Association descriptions in Chapter 5: the  zonal classification and the Ecoregion classification. The former is a climatic classification based on characteristic vegetation on average or zonal terrestrial sites that best reflect the influence of climate, independent of site conditions. Site Association occurrence by zone is outlined in table format at the beginning of each section of Chapter 5. The Ecoregion classification takes a biophysical approach and is a combination of physiography and broad climatic classification. It provides a geographic context for describing Site Association provincial distribution and is used in the general descriptions for each Site Association. Biogeoclimatic () zones Fourteen  zones occur in British Columbia (Figure 2.4). For discus- sions of wetland ecosystem distribution, we have paired similar zones: the BG and PP, the SBS and SBSP, and the BWBS and SWB. The Alpine Tundra (AT) is not covered in this guide. More detailed descriptions of the zones can be found in Meidinger and Pojar (1991). Bunchgrass / Ponderosa Pine (BG/PP) The Bunchgrass and Ponderosa Pine zones are limited to low-elevation areas in the rainshadow of the southern mountains where dry, hot growing-season climates prevail. Grasslands, and ponderosa pine or Douglas-fir forest, are the dominant upland vegetation. Wetlands are primarily marshes. Boreal White and Black Spruce / Spruce – Willow – Birch (BWBS/SWB) The Boreal White and Black Spruce and Spruce – Willow – Birch zones occupy the northern quarter of the province at low to high elevations. These zones have short, cool summers and very long, very cold winters. Upland vegetation is primarily fire-initiated, white and black spruce forests. Peatland formation is favoured in these areas and extensive bogs and fens occur in low-relief landscapes. Coastal Douglas-fir (CDF) The Coastal Douglas-fir zone is restricted to low-elevation (<150 m) coastal areas in the rainshadow of Vancouver Island. This zone has a

Chapter 2 Ecology and classiﬁcation of wetland ecosystems 17  . Biogeoclimatic zones of British Columbia.

Mediterranean climate characterized by warm, dry summers and mild, wet winters. Upland vegetation is primarily Douglas-ﬁr, hemlock, and western redcedar forest. Both peatland and mineral wetlands are common. Coastal Western Hemlock (CWH) The Coastal Western Hemlock zone occurs at low to middle elevations west of the Coast Mountains. This zone has cool, wet summers and mild, wet winters. Natural upland vegetation is primarily old-growth western redcedar, western hemlock, and Sitka spruce forests. This climatic regime favours peatland formation.

18 Wetlands of British Columbia: A Field Guide to Identification Engelmann Spruce – Subalpine Fir (ESSF) The Engelmann Spruce – Subalpine Fir is the uppermost forested zone in the southern three-quarters of the province. The zone has short, cool summers and long, cold, snowy winters. Upland vegetation is closed to patchy forest dominated by Engelmann spruce and subalpine fir. Wet- lands are primarily peatlands. Interior Douglas-fir (IDF) The Interior Douglas-fir zone dominates low to mid elevations of the south-central Interior. This zone has warm, dry summers and cool, dry winters. Upland vegetation is fire-maintained Douglas-fir forest. Wet- lands are diverse in this zone with a mix of peatland and mineral wetland. Interior Cedar – Hemlock (ICH) The Interior Cedar – Hemlock zone occurs on the windward Interior mountains and Coast transition valleys at low to middle elevations. This zone has mild, moist summers and cool, wet winters. Upland vegetation is forest dominated by western redcedar, western hemlock, and interior spruce. Peatlands are favoured but mineral wetlands also occur. Mountain Hemlock (MH) The Mountain Hemlock zone occurs at high elevations along the windward Coast Mountains. This zone has short, cool, wet summers and long, cool, snowy winters. Upland vegetation is open mountain hemlock forest. Wetlands are almost exclusively fens and swamps. Montane Spruce (MS) The Montane Spruce zone occurs at middle elevations in the southern Interior. This zone has short, warm summers and cold winters. Upland vegetation is fire-maintained forest of lodgepole pine and interior spruce. Wetlands are primarily peatlands but mineral wetlands are not uncommon. Sub-Boreal Pine – Spruce / Sub-Boreal Spruce (SBPS/SBS) The Sub-Boreal Pine – Spruce and Sub-Boreal Spruce zones occur in the central Interior at low to middle elevations. These zones have a continental climate with cool, dry to moist, short summers and cold winters. Upland vegetation is fire-maintained forest of interior spruce and lodgepole pine. Peatland formation is favoured in this climate.

Chapter 2 Ecology and classiﬁcation of wetland ecosystems 19 Ecoprovinces Ecoprovinces are used in this guide to describe general distribution of units in British Columbia (Figure 2.5). Northern Boreal Mountains Ecoprovince (western BWBS, SWB) The general character of this Ecoprovince is one of mountains and plateaus separated by wide valleys and lowlands. Short, cool growing seasons promote formation of peatlands in most wet depressions. Con- sequently, subdued terrain may be covered by large expanses of fen and shrub-carr ecosystems. Marshes and swamps are common in association with lake and river systems. Taiga Plains Ecoprovince (eastern BWBS) This Ecoprovince is a large lowland of poorly drained, glaciolacustrine deposits to the east of the northern Rocky Mountains. The climate is subarctic with very cold winters and short cloudy summers. This region has some of the highest concentrations of wetlands in the province (30%

Northern Boreal Taiga Plains Mountains

Boreal

Plains Sub-Boreal

Interior

Coast Central

and Interior Southern Mountains Interior

Southern Mountains

Interior

Georgia Depression

 . Ecoprovinces of British Columbia.

20 Wetlands of British Columbia: A Field Guide to Identiﬁcation areal extent). Bogs predominate, though fens and swamps occur along the sluggish streams that drain the region. Boreal Plains Ecoprovince (eastern BWBS) This Ecoprovince lies east of the Rocky Mountains and consists of low- relief plateaus, plains, prairies, and lowlands. The climate is typically continental since most of the moist Paciﬁc air has dried crossing succes- sive ranges of mountains before it reaches the area. Winters are cold. Bogs are common throughout. Sub-Boreal Interior Ecoprovince (SBS, ESSF) This Ecoprovince lies to the east of the Coast Mountains and consists of low-lying plateaus and several mountain ranges. Western areas are in the rainshadow of the Coast Mountains while eastern areas on the windward side of the Rocky Mountains are wet. Wetlands are primarily fens, with marshes and swamps associated with lakes and streams. Central Interior Ecoprovince (IDF, SBPS, MS, dry ESSF) This Ecoprovince lies in the rainshadow to the east of the Coast Moun- tains and is primarily rolling plateaus. The area has a typical continental climate: cold winters, warm summers, and a precipitation maximum in late spring or early summer. Extensive level terrain harbours a high concentration of wetlands, primarily fens. Southern Interior Ecoprovince (BG, PP, IDF, MS) This Ecoprovince lies in the rainshadow of the Coast and Cascade Mountains and contains some of the warmest and driest areas of the province in summer. Semi-arid conditions limit peat formation except at higher elevations; therefore, marshes and swamps are most common. Some potholes and shallow lakes may experience severe evaporation and drawdown during the summer months, resulting in the accumulation of salts and formation of distinctive ecosystems. Southern Interior Mountains Ecoprovince (ICH, ESSF, BG/PP) This Ecoprovince is the highlands and mountains east of the southern Interior plateaus. There are two distinct climate regimes—wet in the mountains and dry in southern Rocky Mountain Trench. Wetlands are not common in this region, occupying mainly the valley bottoms where many have been altered by development. Coast and Mountains Ecoprovince (CWH, MH) Encompassing the coastal mountain ranges, this Ecoprovince experi- ences a maritime climate characterized by abundant precipitation and mild summer and winter temperatures The geology is predominantly

Chapter 2 Ecology and classiﬁcation of wetland ecosystems 21 granites poor in minerals, with little or no glacial till. Consequently, wetlands are primarily bogs even though most sites receive groundwater seepage. These bogs are extensive on gently to steeply sloping terrain of the outer Coast. Georgia Depression Ecoprovince (CDF, dry CWH) The southeast corner of Vancouver Island, the Gulf Islands, and part of the adjacent mainland experience mild winters, warm summers, and moderate precipitation. Wetlands are not uncommon in this landscape (6% of the area), but most have been modiﬁed by human activities; some have been lost completely. The northern range limit of many plant and animal species is within this region and this, combined with the mild climate, produces ecosystems not found elsewhere in British Co- lumbia.

SUMMARY OF SAMPLING AND ANALYSIS METHODS

The Site Associations presented here were developed using vegetation and environmental data from approximately 2600 plots located throughout British Columbia. The principal data sources include wetland classification projects (900 plots), Biogeoclimatic Ecosystem Classifica- tion program (1000 plots), ecosystem mapping projects (400 plots), and thesis data (300 plots) (Beil 1969; Revel 1972; Annas 1974; Ceska 1978; Banner 1983). Most plot sampling before 1998 used standard methods outlined in Describing Ecosystems in the Field (Luttmerding et al. 1990). Sampling after 1998 followed the Field Manual for Describing Terrestrial Ecosystems (Province of British Columbia 1998). Sites sampled specifically for this guide were selected using available resource information such as topographic maps, air photos, and forest cover maps. Ecosystem plots were located on suitable sites that were homogeneous and relatively undisturbed and the position fixed by . A sample plot size of 20 by 20 m (400 m2) was used for most sites but was reduced or skewed to fit smaller areas such as pocket wetlands or elongated zones within larger complexes. In each plot, vegetation and environmental data were collected on the 882 ecosystem field form according to standard procedures outlined in the Field Manual for De- scribing Terrestrial Ecosystems (Province of British Columbia 1998). Plant species on each site were listed by layer, with an estimate of species percent cover for each layer and for total cover. Unknown specimens were collected and identified.

22 Wetlands of British Columbia: A Field Guide to Identification One soil pit (for mineral soils) or peat core (for organic soils) was established in each plot. Texture, decomposition, depth, and other descriptors where noted for each soil horizon, and the soil type classified according to the Canadian System of Soil Classification (Canada Soil Survey Com- mittee 1998). Humus forms were classified using Green et al. (1993). Peat cores were taken using a modified Hiller peat sampler. We used a combination of ordination and tabular analysis to classify ecosystem plots into Site Associations. We initially subjected the plots for each biogeoclimatic subzone to tabular analysis using Braun- Blanquet methods (Mueller-Dombois and Ellenberg 1974), with the assistance of an ecological database program, 97 ver. 2.0 (MacKenzie and Klassen 1999). Ordination by Detrended Correspondence Analysis () using - ver. 4.0 (McCune and Mefford 1999) was performed and combined with an overlay of the initial tabular classification to aid in assigning plots and differentiating units. Subzone units (Site Se- ries) were combined into the Site Associations appearing in this guide using tabular analysis and ordination of summary values from each subzone unit. Environmental descriptions for each Site Association were produced from summary environmental tables generated from 97 ver. 2.0. Ordinations diagrams of the final classification are presented in Appendix 1.

Chapter 2 Ecology and classification of wetland ecosystems 23 Procedures for 3 site description and identiﬁcation

Site description is the ﬁrst step in the process of classiﬁcation and an understanding of ecosystem function.The measurement and recording of basic attributes of this coastal bog in the Ecstall River valley will give insight into its controlling environmental factors and habitat values. DESCRIBING AND IDENTIFYING SITE UNITS

Site unit identification requires: 1) accurate description of site, soil, and vegetation characteristics 2) use of the various aids and descriptive materials in this guide to determine the site unit that best matches these characteristics. It should not be expected that sites will perfectly match all details in the description of site units in this guide. The descriptions presented here represent a range of conditions around a central concept for a population of more variable individual sites that are part of the Association. It is also important to note that the classification is based on relatively undisturbed wetlands and that identification of highly disturbed or heavily managed wetlands is problematic. This guide describes ecosystems that recur throughout the landscape and appear to be relatively stable in vegetation composition. Although this guide represents the majority of wetland sites in the province it is likely that users will find units that do not match any of the units described. This can be for one of four reasons: 1) the plot location was placed in a transitional area between two ecosystems 2) the site is a hybrid of two types 3) the site is disturbed or in transition from one type to another 4) the site represents a new ecosystem not previously recognized. If the user feels that this is the case, data and information on the site should be forwarded to the Research Branch. This guide is just one tool to help describe and compare wetland and related ecosystems and must be supplemented with practical knowledge, experience, and judgement. Describing sites The following steps are the suggested approach to describe a wetland or wetland-related ecosystem: 1) Select sample area. Plots should be placed in homogeneous areas within wetlands; sites that cross community lines and are heterogeneous are not useful for ecosystem classification. A standard 20 × 20 m plot is ideal; however, wetland communities often occur as narrow bands within a larger wetland. Plots should be made-to-fit in these community types. Microtopographic variation in site and vegetation characteristics is permissible but sites should not include pronounced differences in site, soil, or vegetation.

26 Wetlands of British Columbia: A Field Guide to Identiﬁcation 2) Determine and record site and soil features. The methodology outlined in Province of British Columbia (1998) should be followed. Site features should represent the entire sample area. Soils should be recorded from a soil pit of at least 60 cm depth. A peat core to >160 cm is preferred on peatland sites. For sites with strong microtopographic features, descriptions of hummock and hollow soils are important. Table 3.1 lists some important site and soil attributes that should be described. 3) Determine and record vegetation features. Identify and record percent ground cover of as many plant species as possible. Unknown species, especially those that form a large component of the ground cover, should be collected and preserved for proper identiﬁcation. Species are recorded by layer. • trees: woody plants >10 m • tall shrubs: woody plants >2 m and <10 m • low shrubs: woody plants <2 m • herbaceous species and dwarf shrubs: non-woody species, aquatics, and dwarf shrubs (woody species that are largely <10 cm at maturity) • mosses and lichens: all mosses, liverworts, and lichens 4) Estimate hydrodynamic index, pH, absolute soil moisture, and nutrient regime using tools outlined in Section 5.0.

 . Important site and soil features for identifying site units

Feature Deﬁnition and description  unit From  maps Position in wetland Relationship to other ecosystems Soil texture Soil texture or von Post decomposition of rooting zone Surﬁcial material Mode of deposition of soil parent materials Watertable depth Depth to watertable Humus form Humus form order of the surface soil organic layers Hydrogeomorphic system Hydrological system of site Microtopography Type and strength of surface microtopography

Chapter 3 Procedures for site description and identification 27 Identifying site units Once the site, soil, and vegetation information has been recorded and soil moisture, nutrient, and hydrodynamic index have been determined, the Site Class and Association can be determined. 1) Determine the Class by using the Class key, the summary table of characteristics, and/or the descriptions of Classes presented in Chap- ter 4. The key is dichotomous and presents a series of choices between two alternatives until the name of the appropriate Class is reached. Review the brief Class descriptions following or the one- page descriptions at the start of each Class section. 2) Go to the appropriate Class section of Chapter 5 and review the species importance table for the Associations of the identified Class. Compare the vegetation of the site under consideration with that of the table and select the closest match. Confirm the correct identification by comparing species and site descriptions for the Site Association. Sites intermediate between bogs and fens, fens and swamps, marshes and fens, and swamps and low-bench flood ecosystems should be expected to occur. The user may have to review the Site Associations for more than one Class to correctly identify equivocal sites.

28 Wetlands of British Columbia: A Field Guide to Identiﬁcation Site classes 4

The Class level of classification is useful where site information is not readily available or the scale of application is broad.The ecosystem Classes of this location along the Torpy River, McGregor Mountains, have distinct visual signatures that are readily identifiable from the air. This chapter provides descriptions and tools for the identification of wetland and related ecosystem Site Classes. Three tools can be used to determine Site Class: the key to Site Classes, the summary table of characteristics, or summary descriptions. More detailed descriptions of Classes are found at the beginning of each section in Chapter 5 along with a species importance table of Associations in the Class. The user can use one or all of these tools to identify the Site Class. A suggested approach follows three steps: 1) Use the key to tentatively identify a Class. 2) Review the summary characteristics table (Table 4.1) and short descriptions to substantiate the results from the key. 3) Proceed to the relevant section of Chapter 5 and review Association descriptions.

KEY TO SITE CLASSES

Figure 4.1 is dichotomous key for determination of Site Class. It uses a combination of simple vegetation and site features to tentatively identify the Site Class.

BRIEF DESCRIPTIONS OF SITE CLASSES

The following descriptions outline the essential characteristics of the Site Classes described in this guide. Figure 4.2 shows the distribution of each of the Classes on the modified edatopic grid. Bog Wetland Class (Wb) Bogs are shrubby or treed, nutrient-poor peatlands with distinctive communities of ericaceous shrubs and hummock-forming Sphagnum species adapted to highly acid and oxygen-poor soil conditions. Bogs develop in basins where peat accumulation has raised the wetland surface above groundwater flow, or, less commonly, where groundwater is very low in dissolved nutrients (e.g., flows from granitic parent material). Fen Wetland Class (Wf) Fens are peatlands where groundwater inflow maintains relatively high mineral content within the rooting zone. These sites are characterized by non-ericaceous shrubs, sedges, grasses, reeds, and brown mosses. Fens develop in basins, lake margins, river floodplains, and seepage slopes, where the watertable is usually at or just below the peat surface for most of the growing season.

30 Wetlands of British Columbia: A Field Guide to Identiﬁcation  . Summary of characteristics for wetland and related ecosystem Site Classes

Site Realm/ Cover Group Site Class Environmental features types Species groups

Wetland Wet or Very Wet SMR Realm Bogs +/- ombrotrophic Conifer Sphagnum mosses, pH < 5.5 treed or ericaceous shrubs, > 40 cm ﬁbric/mesic peat low shrub and conifers

Fens Groundwater-fed Graminoid Deciduous shrubs, pH > 5.0 or low sedges, and > 40 cm ﬁbric/mesic peat shrub brown mosses

Marshes Mineral soils or well-humiﬁed peat Graminoid Large emergent sedge, Protracted shallow ﬂooding or forb grass, forb, or horse- (0.1–2.0 m) tail species

Swamps Mineral soils or well-humified peat Tall shrub Conifers, willows, Temporary shallow flooding or alders, forbs, grasses (0.1–1.0 m) forested leafy mosses Significant water flow

Shallow Permanent deep ﬂooding Aquatic Aquatic species waters (0.5–2 m) Emergent vegetation < 10% cover

Estuarine Tidal, brackish water Realm Estuarine High intertidal and supratidal zones Graminoid Grasses, sedges, and meadow Brief semi-diurnal tidal ﬂooding by forbs tolerant of di- Class brackish water urnal ﬂooding and brackish water

Estuarine Intertidal Graminoid Salt-tolerant emergent marsh Diurnal tidal ﬂooding by salt water or forb graminoids and suc- Class culents

Flood Riparian flood zone Group of High Benches above normal waterflow Coniferous Upland species of Terrestrial bench Brief flood period forested seepage sites Realm

Mid Elevated benches ﬂooded most Deciduous Flood-tolerant decid- bench years for < 21 days treed or uous trees and Areas of sedimentation forested shrubs

Low Site directly adjacent to watercourse Tall Flood-tolerant shrubs bench Annual ﬂood >21 days deciduous Signiﬁcant annual erosion and shrub deposition

Transition “Special” factor Classes from Shrub- Frost-prone depressions with ﬁne- Low shrub Deciduous low Terrestrial carr to medium-textured moist soils shrubs, grasses, and Realm forbs

Saline Semi-arid climate Graminoid Flood and salt- meadow Slightly to highly saline soils tolerant graminoids Brief periods of inundation and forbs

Chapter 4 Site classes 31 1 Sites tidal and influenced by salt water ...... Estuarine (Section 5.6) 1a Sites non-tidal or in freshwater tidal reaches of river...... go to 2 2 Wet or Very Wet sites. Soils Gleysols or Organics...... go to 3 2a Soil moisture regime hygric or drier. Soil types variable ...... go to 8 Wetland Ecosystems 3Sites permanently flooded and with < 10% emergent vegetation go . to .4 ...... 3a Sites with >10% emergent cover ...... go to 5 4 Sites with > 10% cover of rooted aquatic plants ...... Shallow-waters (Section 5.5) 4a Sites with < 10% cover of rooted aquatic plants ...... Unclassiﬁed aquatic ecosystem 5 Sites with mineral soil or a surface tier of humic peat (von Post 7 or greater) ...... go to 6 5a Sites with organic soils and surface tier dominated by fibric or mesic sedge or moss peat ...... go to 7 Mineral Group 6 Sites dominated by tall shrubs or trees (> 10% cover) ...... Swamps (Section 5.4) 6a Sites dominated by emergent grass-like species. Shrub and tree cover < 10% ...... Marshes (Section 5.3) Peatland Group 7 Sphagnum Groups I & III dominate moss layer. Ericaceous spp. common. Peat pH < 5.5. . Bogs (Section 5.1) 7a Bryophyte layer not dominated by Sphagnum. Graminoids dominant. Peat pH > 5.0 . . . Fens (Section 5.2) Terrestrial Ecosystems 8 Sites not inundated by floodwaters. Soils non-cumulic ...... go to 9 8a Sites periodically flooded by adjacent rivers or lakes. Soils commonly layered (cumulic) ...... go to 10 9 Sites not affected by special site factors such as severe cold-air ponding, ...... Sites described by BEC or soil salinity. “Normal” forest or grassland communities 9a Sites affected by special site factors such as severe cold-air ponding, or soil salinity ...... go to 13 Flood Group 10 Tree cover < 10%. Always immediately adjacent to and not much ...... go to 11 elevated above waterbodies 10a Tree cover > 10%. May be adjacent to active channel or at some distance from open ...... go to 12 water on elevated benches and terraces. Soils with some horizon development 11 Sites with continuous shrub cover; flooded for moderate periods ...... Low benches (Section 5.7) 11a Sites with sparse shrub cover (< 10%). Flooded for prolonged periods . . . Active channel (not described) 12 Coniferous trees usually predominate. Understorey vegetation resembles . . . . . High benches (BEC units) upland seepage sites 12a Deciduous trees predominate; conifers limited to elevated microsites . . . . . Middle benches (Section 5.7) Other Terrestrial Classes 13Sites with > 10% low shrubs in frost hollows or gradualShrub-carrs slopes (Sectionexperiencing 5.8) . . . . cold-air drainage 13a Sites dominated by graminoids and salt-tolerant forbs. In drawdown ...... Saline meadows (Section 5.8) zone of ephemeral ponds or shallow lakes. Mainly in subzones with extensive natural grasslands (e.g., IDF, SBPS, BG, PP)

 . Dichotomous key for the identification of Site Class.

32 Wetlands of British Columbia: A Field Guide to Identification Marsh Wetland Class (Wm) Soil Nutrient Regime A marsh is a shallowly flooded ABCD E F Terrestrial Ecosystems Meadows mineral wetland dominated by M Low and Shrub-carrs middle emergent grass-like vegetation. benches Estuary VM Meadow A fluctuating watertable is typical Swamps Estuary in marshes, with early-season high W Bogs Marsh watertables dropping through the AkVA N MA SA Marshes growing season. Exposure of the VW substrate in late season or during Soil Moisture Regime Fens dry years is common. The sub- pH strate is usually mineral, but may St Sl Mo Dy VD have a well-decomposed organic Hydrodynamic Index veneer derived primarily from  . Site Class distribution on the marsh emergents. Nutrient avail- modified edatopic grid. Shallow-water wetlands do ability is high (eutrophic to not fit this conceptual model and are not indicated. The wetland edatopic grid is described in detail in hyper-eutrophic) due to circum- Chapter 5.0. neutral pH, water movement, and aeration of the substrate. Swamp Wetland Class (Ws) A swamp is a forested, treed, or tall-shrub, mineral wetland dominated by trees and broadleaf shrubs on sites with a flowing or fluctuating, semipermanent, near-surface watertable. Tall-shrub swamps are dense thickets, while forested swamps have large trees occurring on elevated microsites and lower cover of tall deciduous shrubs. Both types of swamps have abundant available nutrients from groundwater and often have surface standing water. Swamps may be underlain with peat but this is well decomposed, woody, and dark. Shallow-water (Aquatic) Wetland Class (Wa) Aquatic wetlands are shallow waters dominated by rooted, submerged and floating aquatic plants. These communities are always associated with permanent still or slow-moving waterbodies such as shallow potholes or deeper ponds and lakes. Shallow-water sites are usually permanently flooded; rarely they may become exposed during extreme drought years. Shallow-water communities most commonly occur where standing water is less than 2 m deep in midsummer. Aquatic plants may root in mineral soils or in well-humified sedimentary peat. Saline meadow Transition Class (Gs) Saline meadows are grass-, rush-, or halophyte-dominated sites that occur on periodically saturated and occasionally inundated sites, where

Chapter 4 Site classes 33 watertable decline is caused mainly by evaporation and where salts accumulate. These conditions occur only in dry climates. After a brief period of inundation, the watertable drops below the rooting zone during most of the growing season, resulting in a well-aerated rooting medium. These ecosystems are part of a Grassland Group of terrestrial ecosystems. Shrub-carr Transition Class (Sc) A shrub-carr is a shrub-dominated ecosystem that develops on frost- prone sites with moist or very moist soils. These sites are seasonally saturated but rarely inundated (see flood ecosystems) and may have watertables perched at depth. Shrub-carrs frequently border wetlands or occur in frost-prone hollows in cold and dry climatic regions. A strongly mounded soil surface is typical, and shrubs of 1–2 m occur mainly on these elevated microsites. These ecosystems are part of a Shrubland Group of terrestrial ecosystems. Low bench Flood Class (Fl) Low bench ecosystems occur on sites that are flooded for moderate periods (< 40 days) of the growing season, conditions that limit the canopy to tall shrubs, especially willows and alders. Annual erosion and deposition of sediment generally limit understorey and humus development. Middle bench Flood Class (Fm) Middle bench ecosystems occur on sites briefly flooded (10–25 days) during freshet, allowing tree growth but limiting tree species to only flood-tolerant broadleaf species such as black cottonwood and red alder. High bench Flood Class (Fh) High bench ecosystems occur where flooding rivers produce lengthy subsurface flow in the rooting zone but only periodic, brief inundation. Surface flooding may occur from as frequently as several times annually to only during extreme flood years. These periods of flooding are generally not restrictive of plant species; plant communities are similar to adjacent upland forests on seepage sites. High bench Site Series are described in  field guides and are not presented in this guide. Estuarine Marsh Class (Em) An estuarine marsh is an intertidal ecosystem that is flooded diurnally and has simple communities dominated by salt-tolerant emergent graminoids and succulents. These marshes occur in the middle to upper tidal zones of estuaries where saltwater influences predominate.

34 Wetlands of British Columbia: A Field Guide to Identiﬁcation Estuarine meadow Class (Ed) Estuarine meadows occur in the high intertidal and supratidal zones of estuaries, where tidal ﬂooding occurs less frequently than daily and is tempered by freshwater mixing. Species composition is relatively diverse, typically with a mix of graminoids and forbs.

Polygonum amphibium, water smartweed

Chapter 4 Site classes 35 Site associations 5

Classiﬁcation is a process of distilling complex ecosystem variability into simpler, ecologically meaningful units. Many apparently unique sites, such as this visually distinctive Chamisso's cotton-grass fen, may actually be very similar to others in overall species composition and environment. This chapter presents detailed Site Class and Association descriptions in standardized format. The chapter has eight sections, one for each Class or Group. Each section includes a four-page Class description followed by one-page Site Association fact sheets. Associations for which there is little current information or that are similar to more common Associa- tions are given a single paragraph outlining the characteristics of such units at the end of each Class section.

LAYOUT AND CONVENTIONS FOR SITE CLASS DESCRIPTIONS

A standard-format, four-page Class description precedes one-page fact sheets for each of the Site Associations in the Class. Definition: A concise definition of the Class. Edatopic Grid: General location of the Class on the edatopic grid. Vegetation: A description of the vegetation structure and composition typical and characteristic of the Class. Landscape: Provincial distribution and locations in the landscape where the Class is found. Hydrology and Soils: Soil types and hydrological characteristics that are typical for the Class. Other Comments: Additional information. Conservation Issues: Common Class characteristics of importance to management and conservation of ecological function. Distribution of Site Associations by  Zone: This presents a list of Associations described in the guide and their distribution by biogeoclimatic zone. The abundance of the type relative to other wetland types in the zone is given a ranking: x = incidental/rare: occurs infrequently in the zone, <5% of wetland occurences. xx = minor/uncommon: 5–25% of wetland occurences. xxx = major/common: >25% of wetland occurrences, often occupies extensive areas. In some cases, a superscript letter is used to indicate a specific part of the zone where the unit occurs. Species importance table Species that are distinctive for at least one of the Associations are included in the species importance table and indicated by relative importance:

38 Wetlands of British Columbia: A Field Guide to Identiﬁcation ❘ Infrequent: occurs very sporadically in the unit. Usually <30% of plots. ❘❙ Uncommon: occurs on a minority of sites (<30%) and with low cover. ❘❙❚ Common: occurs on many sites (30–60%), often low cover but occasionally with appreciable cover (to 10%). ❘❙❚❚ Abundant: occurs on most sites (>60%), occasionally prominent on some sites (± 10%). ❘❙❚❚❚ Very abundant: occurs on nearly all sites (>80%); prominent species (± 10%). ❘❙❚❚❚❚ Dominant: occurs on all sites (>95%); the most abundant species on most sites (>25% cover).

SITE ASSOCIATION FACT SHEET

Site Association fact sheets (Figure 5.0.1) describe each unit as a range or a summary of average conditions. Fact sheets seldom describe the precise conditions of a given plot, but provide the “central concept” for the unit and give insights into the typical conditions that can be expected within a Site Association. Name (1): Name of Site Association deﬁned by diagnostic or leading plant species. Common names are placed in the header and scientiﬁc names below. General Description (2): Brief description of distribution, landscape position, hydrology, soils, and vegetation. Characteristic Vegetation (3): Plant species commonly found in the Site Association, grouped by cover layer. Species importance is coded by font: • Dominant species appear as italicized, bold, and underlined • Abundant and very abundant species as italicized and bold • Common species as italics Comments (4): Additional information on the Association including successional relationships, associated ecosystems, and similar previously described units. Photograph (5): Photo of a typical example of the Site Association. Wetland Edatopic Grid (6): The edatopic grid depicts the location of the

Chapter 5 Site associations 39 Lodgepole pine – Bog rosemary – Peat-moss Wb02 1 Pinus contorta – Andromeda polifolia – Sphagnum

General Description 2 7 Lodgepole pine – Bog rosemary – Peat-moss bogs are scattered throughout the wet regions of the Central and Sub- Boreal Interior (ICHvk/wk and SBS vk/wk) at elevations below 1100 m. They occur in closed basins, isolated zones in larger peatlands, and occasionally around acidic peatland lakes. Pinus contorta usually dominates in the sparse and stunted canopy but Picea mariana, Tsuga heterophylla, or Abies lasiocarpa can form a 5 significant component. Dwarf woody plants such as Kalmia microphylla and Andromeda polifolia are prominent. On some sites Carex pauciflora and Eriophorum angustifolium may also be prevalent. Soils are Fibrisols and Mesisols typically composed of deep Sphagnum peat but sometimes they occur on peat veneers over dense, fine-textured glaciolacustrine deposits. 3 Characteristic Vegetation Wetland Edatopic Grid 6 (0 - - 40) Tree layer 5 Soil Nutrient Regime Picea mariana, Pinus contorta ABCD E F Shrub layer (6 - 32 - 95) Betula nana,Ledum groenlandicum, VM Picea mariana, Pinus contorta Herb layer (3 - 30 - 95) W Andromeda polifolia,Carex pauciflora, VA MA D Eriophorum angustifolium,Kalmia micro- V x phylla,Oxycoccus oxycoccos, Rubus VW e chamaemorus SA Ind

Soil Moisture Regime Mo Dy p mic Moss layer (35 - 95 - 100) H N l Group I S Pleurozium schreberi,Sphagnum Ak St Hydrodyna 4 Comments The Wb02 Site Association is one of several Bog Site Associations that occur in the interior rainforest.The Wb02 likely reﬂects the “true bog”or “climax”condition of long-term peatland succession in the cool and warm subzones of the interior rainforest of the Central and Sub- Boreal Interior.There are hemlock-dominated bogs (Wb04) in colder and snowier subzones and black spruce–dominated bogs in areas with more boreal climates (such as frost pockets). The Wb07 may be the successional precursor to the Wb02. The Wb02 includes Site Series ICHmm/07, ICHvk2/07,SBSvk/08,and SBSwk3/05 but occurs more widely in the wet SBS and ICH.

Chapter 5.1 Bogs 57

 .. Example Site Association fact sheet.

Site Association in relation to important environmental features. The shaded area represents the estimated environmental range of the Site Association in relation to actual soil moisture and nutrient regime, acidity/alkalinity, and hydrodynamic index. A description of the wetland edatopic grid is found below. Hydrogeomorphic Icon (7): Symbol(s) indicating in which hydrogeomorphic group the unit commonly occurs. The hydrogeomorphic classiﬁcation describes the topographic position and hydrology of sites (MacKenzie and Banner 2001).

40 Wetlands of British Columbia: A Field Guide to Identification Estuarine System: Sites at the confluence of fluvial and marine environments affected by tides

Fluvial System: Sites associated with flowing water and subject to flooding, erosion, and sedimentation Lacustrine System: Sites at lakeside, directly affected by lake hydrological processes (e.g., wave action, flooding, and sedimentation) Palustrine System, Sites in depressions and other topograph- Basins and ic low points with the watertable near or Hollows: at the surface; receive water mainly from groundwater and precipitation Palustrine System, Sites associated with small waterbodies Ponds and Potholes:

Palustrine System, Sloping sites with near-surface ground- Seepage slopes: water seepage

The Wetland Edatopic Grid The edatopic grid used in this guide is a modification of the model used in . The  grid uses soil moisture and nutrient regimes as major site descriptors for comparing forested ecosystems. However, additional factors are important in wetland and riparian ecosystems, including acidity/alkalinity (as a correlate to availability of base cations) and magnitude of lateral flow or vertical fluctuation (hydrodynamics). In the Wet and Very Wet portions of the edatopic grid, two tangent environmental axes have been added to accommodate these important site factors, based on concepts presented in Vitt (1994). The grid should be viewed as a conceptual model; it has not been rigorously tested with field data. Several edatopic grid overlays of vegetation characteristics to assist in in- terpreting the grid are presented in Appendix 2. The four axes of the grid (Figure 5.0.2) describe specific site characteristics and are defined as follows: Actual Soil Moisture Regime () is the average amount of soil water annually available for evapotranspiration by vascular plants over

Chapter 5 Site associations 41 several years (Pojar et al. 1987). Soil Nutrient Regime There are nine moisture categories ABCD E F from Very Dry to Very Wet. Wet- M lands are found only on Wet to VM Very Wet sites. Related ecosystem classes are also found on Moist W and Very Moist sites. The wetland AkVA N MA SA edatopic grid is therefore limited VW to this range. The definitions for Soil Moisture Regime pH soil moisture categories used in the guide are defined as: St Sl Mo Dy VD Hydrodynamic Index Moist (): No water deficit  .. A modified edatopic grid occurs. Current need for showing pH and hydrodynamic index axes for Wet water does not exceed supply; and Very Wet sites. temporary groundwater table may be present. Unless otherwise limited, supports forest. Very Moist (): Rooting-zone groundwater present during the growing season (water supply exceeds demand). Groundwater table > 30 cm below the surface. Unless otherwise limited, supports forest. Wet (): Rooting-zone groundwater present during the growing season (water supply exceeds demand). Groundwater table between 0 and 30 cm below the surface. Can support tall shrubs and trees. Very Wet (): Groundwater table at or above the ground surface during the growing season. Will not support tall shrubs or trees but can support low shrubs. Soil Nutrient Regime () is the essential soil nutrients available to vascular plants over a period of several years (Pojar et al. 1987). Six  classes are recognized from Very Poor to Alkaline/Saline. Wetland and wetland-related ecosystems can occur throughout the range. Some indicators of  in wetlands are presented in Figure 5.0.3. pH (acidity/alkalinity) is a correlate measure of base cation availability. This is primarily of importance for peatlands and less important for hydrologically dynamic systems. Five categories are recognized from Very Acid to Alkaline. Generally, as acidity increases, available base cations decrease, resulting in reduced site productivity. Very Acid (): (<4.5 pH) sites are true bogs with high cover of Sphagnum Group I or III mosses and few minerotrophic indicators.

42 Wetlands of British Columbia: A Field Guide to Identiﬁcation (alkaline) excess alkali excess very clear blue-green and blue-green Marl Low Mineral continuous seepage continuous Saprimoder green-brown and turbid green-brown dark diurnal exposure of substrate diurnal exposure seasonal seepage seasonal exposure of substrate seasonal exposure Mesimor Medium green-brown and clear green-brown pale Fibrimor stagnant always saturated saturated always High tea colored;tea and turbid brown yellowish-deep A B C DC A B E F <5.0 4.5 – 6.0 5.0 – 6.5 6.0 – 7.4 6.5 – 8.0 8.0+ 1 – 3 3 – 6 4 – 7 7 – 10 8 – 10 very low low average plentiful abundant or salt accumulation SNRAvailable nutrients pH Water of vonPost Poor Very tier surface - Ground ﬂow water site through PoorC:N ratio Surface tier material MediumWater colour Rich Colour of surface peat Surface tier Rich Very saturation Hyper

 .. Environmental characteristics useful for determining nutrient status in wetlands.

Chapter 5 Site associations 43 Moderately Acid (): (4.5–5.5 pH) sites still have high Sphagnum cover but minerotrophic indicators also occur. Peatland sites are considered bogs in this guide but would be poor fens or poor swamps using a “classic” definition. Slightly Acid (): (5.5–6.5 pH) sites are fens or swamps. Tomen- thypnum, Warnstorfii, and Drepanocladus brown mosses are typical for sites with a stagnant or sluggish hydrodynamic index. Neutral (): (6.5–7.4 pH) sites are fens, swamps, or marshes. Species are often a combination of species found on slightly acid and alkali sites. Alkaline (k): (>7.4 pH) sites are dominated by minerophilic bryophytes such as Scorpidium or Campylium mosses on peatland sites. Alkali-tolerant species occur in marshes. The Hydrodynamic Index () has five categories that describe the magnitude of vertical and lateral water movements in the soil on Wet and Very Wet sites. Stagnant (t): Stagnant to very gradually moving soil water. Vertical fluctuations minimal. Permanent surface saturation but minimal or no surface flooding. Basins or hollows with stable water regimes. Abundant organic matter accumulation and high bryophyte cover. Sluggish (l): Gradual groundwater movement through peat or fine-textured mineral soils along a hydrological gradient. Minor vertical watertable fluctuations. Semipermanent soil saturation with some elevated microsites or brief periods of surface aeration. Hol- lows, slopes, and water tracks in basins or lake flats not directly influenced by the waterbody. Abundant peat accumulation and bryophyte cover. Mobile (o): Distinct flooding and drawdown or pronounced lateral water movements. Peripheral areas of peatlands, sites adjacent to open water tracks, small rivulets or ponds, small potholes with relatively stable water regimes, protected lake embayments, or backmarshes in estuaries. Can have deep but well-decomposed accumulations of peat. Patchy bryophyte cover. Dynamic (y): Significant lateral flow and/or strong vertical watertable fluctuations through mineral soils. Potholes in arid climates that experience significant drawdown, wave-exposed shores, floodplain back channels, and protected estuary sites. Little organic accumulation, few bryophytes.

44 Wetlands of British Columbia: A Field Guide to Identification Very Dynamic (): Highly dynamic surface water regime. Ex- posed tidal sites, shallow potholes in arid climates that experience significant drawdown, wave-exposed shores, and sites directly adjacent to and influenced by river flow. No organic accumulation or bryophytes. Estuarine tidal diagram Estuarine ecosystems occur within the intertidal zone, which is defined as the area between chart datum (zero tide on marine charts) and the limit of higher high tides. These ecosystems have Wet or Very Wet soil moisture regimes but the modified edatopic grid used for wetlands does not describe these estuarine ecosystems well. The salinity of floodwaters and the duration of daily tidal flooding are of primary importance for these systems. Therefore, for the Site Associations in Section 5.5, a simple diagram of salinity and relative height above chart datum replaces the wetland edatopic grid (Figure 5.0.4). Salinity of estuarine sites is variable depending on season and magnitude of tide. The highest salinity that these ecosystems experience during the growing season is likely to be the important attribute rather than the average annual or daily salinity. The highest growing-season salinity is described in a six-category system (from Cowardin et al. 1978). Fresh: <0.5 parts per thousand (ppt) salts Oligosaline: Weakly brackish; 0.5–5 ppt salts Mesosaline: Moderately brackish; 5–18 ppt salts Polysaline: Strongly brackish 18–30 ppt salts Eusaline: Normal seawater; 30–40 Salinity Fresh Oligo- Meso- Poly- Eu- Hyper- ppt salts Hypersaline: >40 ppt salts Estuarine Meadows Elevation above chart datum is a Upper corollary for duration of flooding but is dependent on the magnitude Estuarine Marshes of tides in a particular area. The Middle Wetland – Marshes

Tidal Zone Tidal deﬁnitions below are from Howes et al. (1999). Tidal Flats

Lower The Upper Intertidal describes the upper third of the elevation range  .. Distribution of Site Classes between the highest high tide and relative to intertidal zone and salinity. zero tide for a particular area

Chapter 5 Site associations 45 (e.g., 4.07–6.1 m near Prince Rupert but 2.15–3.2 m at Fulford Harbour). These sites are flooded for 5–50% of each tidal cycle. The Middle Intertidal describes the middle third of the elevation range between highest high tide and zero tide (e.g., 2.03–4.07 m for Prince Ru- pert and 1.07–2.15 m for Fulford Harbour). The lower edge of this zone roughly corresponds to the lower vegetation limit. Sites are flooded for 50–90% of the tidal cycle. The Lower Intertidal describes the lower third of the elevation range between highest high tide and zero tide (e.g., 0–2.03 m for Prince Rupert and 0–1.07 m for Fulford Harbour). Sites are flooded for over 90% of the tidal cycle. Sites are primarily unvegetated tidal flats.

TAXONOMIC CONSIDERATIONS

Taxonomy for vascular plants follows the Illustrated Flora for British Co- lumbia (Douglas et al. 1998a, b, 1999a, b, 2000, 2001a, b). Moss taxonomy follows Anderson (1990) and Anderson et al. (1990). Hepatic scientific names and authorities are based on Stotler and Crandall-Stotler (1977). Lichen scientific names and authorities are consistent with Esslinger and Egan (1995). Species equivalents In some cases, a genus or species name is used to represent more than one species or genus. This has been done where taxa have similar ecological requirements, are difficult to distinguish and are likely to be confused by many field workers, or have a contentious taxonomy. Betula nana includes B. pumila Carex limosa includes C. magellanica Equisetum arvense includes E. pratense Equisetum fluviatile includes E. palustre Picea X includes P. glauca, P. engelmannii, and all hybrids Populus balsamifera includes P. balsamifera ssp. balsamifera, and P. balsamifera ssp. trichocarpa Salix barclayi includes S. pseudomonticola Schoenoplectus acutus includes S. tabernaemontani

46 Wetlands of British Columbia: A Field Guide to Identification Sphagnum Group I are widespread peat-mosses of poor sites and include S. angustifolium, S. capillifolium, S. fuscum, and S. magellanicum Sphagnum Group II are primarily peat-mosses of interior intermediate sites and include S. centrale, S. squarrosum, S. subnitens, S. subsecundum, S. teres, and S. warnstorfii Sphagnum Group III are primarily coastal peat-mosses of raised sites and includes S. austinii, S. papillosum, and S. rubellum. Sphagnum Group IV are primarily peat-mosses in water or saturated lawns and include S. cuspidatum, S. lindbergii, S. mendocinum, and S. tenellum Peat-moss groups are based on several sources (Sims and Baldwin 1996; Gignac et al. 1991; Vitt 1994; Belland and Vitt unpublished). Recent taxonomic changes There have been a number of recent taxonomic name changes to common species in the flora of British Columbia. The species and genera following have been changed in this guide to reflect current standands (Meidinger 2002). Alnus tenuifolia and A. viridis have reverted to A. incana and A. crispa, respectively Betula glandulosa is now B. nana The moss genus Drepanocladus has been divided into Drepanocladus, Homatocaulis, Sanionia, Scorpidium, and Warnstorfia genera Potentilla palustris is now Comarum palustris The genus Scirpus has been divided into four genera: Scirpus, Bol- boshoenus, Schoenoplectus, and Amphiscirpus Scirpus acutus and S. validus are now Schoenoplectus acutus and S. tabernaemontani, respectively Scirpus maritimus is now Bolboshoenus maritimus Tofieldia is now Triantha Utricularia vulgaris is now U. macrorhiza

Chapter 5 Site associations 47 5.1 BOGS

1 Classic boreal black spruce bog, Alaska Highway, Boreal Plains (BWBSmw1) 2 A Shore sedge – Peat-moss floating bog, White River near Meziadin Junction (ICHvc) 3 Blanket mire complex, near Prince Rupert (CWHvh2)

Chapter 5.1 Bogs 49 BOG WETLAND CLASS

Deﬁnition A bog is a nutrient-poor, Sphagnum-dominated peatland ecosystem in which the rooting zone is isolated from mineral-enriched groundwater, soils are acidic, and few minerotrophic plant species occur. General Description Vegetation Table 5.1.2 lists the species that are common in Bog Site Associations described in this guide. Bogs are characterized by an abundance of Sphagnum mosses and evergreen woody vegetation (conifers and ericaceous shrubs) adapted to nutrient-poor site conditions. Sphagnum mosses generally drive these systems because they trap base cations, causing the organic soils to acidify and to retain moisture, thus slowing the decomposition rate and promoting peat accumulation. Bogs commonly support stunted coniferous trees that, on true bogs, rarely reach 7 m in height but can reach 15 m on more productive sites. Sparse shrub and herb layers are common. In wetter bogs, where the watertable is at the surface, tree species do not survive and dwarf shrubs are prominent. Interior bog vegetation is similar to that found throughout boreal regions worldwide. However, bogs of the outer Coast have distinctive, globally unusual vegetation. A hypermaritime climate (with moderate annual temperatures, high precipitation, and high ambient humidity) combined with mineral-poor bedrock promotes widespread bog formation on level and sloping terrain. This blanket mire complex is a combination of open, shrubby, and woodland bog types. Landscape Position and Distribution Bogs occur primarily in closed basins, on the periphery of larger peatlands, or occasionally as raised domes in fens. They are common in climatic regions with cool summer temperatures where evapotranspiration rates are low and saturated conditions are maintained throughout the growing season (Table 5.1.1). Extensive bogs occur in the outer coastal lowlands, where precipitation is high and nutrient-poor parent material is common, and in the Taiga Plains, where there is extensive low-relief terrain on impermeable glaciolacustrine deposits. Topogenous bogs are also common in subdued terrain of the sub-boreal and boreal forests.

50 Wetlands of British Columbia: A Field Guide to Identification Hydrology and Soils Bogs develop in basins where Soil Nutrient Regime peat accumulation has raised the ABCD E F surface peat above the watertable, M or, less commonly, where VM groundwater is near the surface but is very low in dissolved min- W Bogs erals and nutrients (Figure 5.1.1). AkVA N MA SA While the groundwater table can VW be well below the soil surface, the Soil Moisture Regime pH upper tier remains saturated throughout the growing season St Sl Mo Dy VD Hydrodynamic Index through the capillary action of living and dead Sphagnum moss-  .. Position of bogs on the es. Bogs are never flooded. edatopic grid. Soils are usually deep peat deposits with at least the upper layers poorly decomposed and derived from Sphagnum moss. Other Comments The traditional definition of bog describes peatland ecosystems that are ombrotrophic (i.e., isolated from groundwater). However, many peatland ecosystems in British Columbia with bog-like vegetation and abundant Sphagnum experience some groundwater contact, especially in microtopographic hollows. This guide includes these ecosystems (variously referred to as poor fens or poor swamps) in the bog wetland class (see Bridgham et al. 1996). Conservation Issues Many typical bog species are not tolerant of flooding and are out- competed by minerotrophic species when nutrient availability is even merely moderate. Therefore, land uses that increase water inputs to these sites can convert bogs to swamp or fen communities. Road construction that diverts runoff into or blocks drainage from bogs is the most common anthropogenic disturbance in non-urban areas. Additions of nitrogen-rich water (such as from sewage or cattle yard runoff) can quickly degrade bog ecosystems by increasing peat decomposition and facilitating invasion of marsh species such as cattail. Removal of water from bogs will lead to an increase in tree growth and cover of upland species and a loss of obligate hydrophytes, but, because of the moisture- holding capacity of Sphagnum peat, many bog species can persist.

Chapter 5.1 Bogs 51 However, Sphagnum is not tolerant of shading, so continued increase in tree or shrub cover generally leads to a decline in peat-mosses. Bog vegetation is generally very slow growing and therefore provides few forage values for larger wildlife. However, the unique combination of plant species and habitat structure supports distinctive arthropod communities. Bogs in the boreal forest and outer Coast are important Sandhill Crane nesting areas. Timber and range values in bogs are essentially nil. More productive bogs have some sizeable trees but regeneration issues are considerable. Peat cutting has occurred in some regions of the province but is not widespread, in part because peat composition in most regions is of poor quality for horticultural uses or fuel.

Empetrum nigrum, crowberry

52 Wetlands of British Columbia: A Field Guide to Identiﬁcation oc oc s xx xxx xxx xx w w x n xxxxx xxxx BGPP BWBS SWB ESSF ICH IDF MS SBS CDF CWH SBPS MH Distribution of Bog Site Associations by biogeoclimatic zone Black spruce – Creeping-snowberryBlack spruce – Peat-moss Lodgepole pine – Bog rosemary – Peat-moss – LingonberryBlack spruce – Peat-moss hemlock – Cloudberry – Peat-mossWestern sedge – Peat-mossWater – Black spruce moss sedge – Fen Water – Tamarack sedge – Peat-moss Water Lodgepole pine – sedge – Peat-moss – Soft-leaved Black spruce horsetail – Peat-moss – Common Black spruce sedge – Peat-mossLodgepole pine – Few-ﬂowered – Peat-moss – Buckbean Black spruce – Peat-mossScheuchzeria – Peat-moss sedge – Buckbean Shore – Peat-moss Labrador – Bog-laurel tea peat-moss pine – CrowberryShore – Tough xxx moss clubrush – Rock juniper – Tufted Common clubrush – Tufted Yellow-cedar pine – Shore xx x xx xxx x x x x x x xxx x x x x x x x x x x x s = southern subzones only x = incidental; < 5% of wetlands xx = minor; 5–25% of wetlands xxx = major; >25% of wetlands w = wet/very wet subzones only n = northern subzones only oc = outer coast (hypermaritime) only Wb01 Wb02 Wb03 Wb04 Wb05 Wb06 Wb07 Wb08 Wb09 Wb10 Wb11 Wb12 Wb13 Wb50 Wb51 Wb52 Wb53  .. 

Chapter 5.1 Bogs 53  .. Bog Species Importance Table

Species Wb01 Wb02 Wb03 Wb04 Wb05 Wb06 Wb07 Wb08 Trees Picea mariana xyzzz xyzz xyzzzz xy xyzzz xyzz x xyzzz Larix laricina xyz xyzzz x Tsuga heterophylla xy xyzzz x Pinus contorta var. latifolia x xyzz xy xy x xyzz x Picea X x x x xyz xy xy xyzz xyz Thuja plicata xx x Pinus contorta var. contorta Chamaecyparis nootkatensis Shrubs Ledum groenlandicum xyzzzz xyzzz xyzzz xyz xyzz xyzz xyzz xyzz Betula nana xy xyz xy x xyzzz xyzzz xyzz xyzz Salix myrtillifolia x x xyz x Lonicera involucrata x x x xy xyz xy Salix pedicellaris x x xy xyz x xy Myrica gale Vaccinium uliginosum x Juniperus communis Herbs Oxycoccus oxycoccos xyz xyz xy xyz xyz xy xyz xy and Gaultheria hispidula xyzz xy x x xy x xy x Dwarf Vaccinium vitis-idaea xyzz x xyz x Shrubs Rubus chamaemorus x xyz xyz xyzzz x xy x x Carex aquatilis/sitchensis xy xy x xy xyzz xyzzz xyzzz xyz Carex disperma x x x x xy x x xyzzz Carex tenuiflora x x xyz Comarum palustre x x x xyz xy xy xyzz Equisetum arvense xy x xyz xy x xyz xyz Carex pauciflora x xyz xy x x x x Andromeda polifolia xyz x x xy Empetrum nigrum xy x xyz x xy xyz x Carex limosa xx xxyx x xy Menyanthes trifoliata xxxxxyx Eriophorum angustifolium xyz xyz x x x Kalmia microphylla xyz xyz x x Scheuchzeria palustris x Drosera anglica Drosera rotundifolia xxxyx xy Coptis trifolia xxy Carex pluriflora x Fauria crista-galli Carex livida x Sanguisorba officinalis Triantha glutinosa xx Trichophorum cespitosum xx Rhynchospora alba Agrostis aequivalvis Lichens Sphagnum Group I xyzzzz xyzzzz xyzz xyzzzz xyzzz xyzzz xyzzzz xyzz and Pleurozium schreberi xyzz xyz xyzzz xyzzz xyz x xyz xyz Mosses Hylocomium splendens xy xyzzz xy x xyz Aulacomnium palustre xyz x xy x xyz xyzzz xyzz xyz Tomentypnum nitens xy x xyzz xyzzz xyz xyzz Sphagnum Group III xxyx xyx x Cladina spp. xy x xyz xy x x Cladonia spp. xy x xy x x Sphagnum Group IV Racomitrium lanuginosum Siphula ceratites Campylopus atrovirens

54 Wetlands of British Columbia: A Field Guide to Identification Wb09 Wb10 Wb11 Wb12 Wb13 Wb50 Wb51 Wb52 Wb53 Common Name xyzzz xy xyzzz x black spruce x tamarack x x xy xy x x western hemlock xyzz xyz xy lodgepole pine xy xy x x spruce x x x xyz xy xyz western redcedar xy xyzz xyzz xyzzz shore pine xyz xyz xyzzzz yellow-cedar xyzz xyz xyzz xy x xyzzz xyzz xyz xyzz Labrador tea xyz xyz xyzz xy scrub birch xy x bilberry willow xy x black twinberry x xyxyxyzx bog willow x xyzz xyz xyzz x sweet gale x xy xyz xy bog blueberry xyz xyzzz xy common juniper xy xyz xyzz xyz xy xyzz xy xy x bog cranberry xy x xy creeping-snowberry xy xxx lingonberry xxyxxxxxyzzxx cloudberry xy xyz xyz x xy xy xy x water sedge/Sitka sedge xy x soft-leaved sedge xxxyx sparse-leaved sedge x x xyz xy xy marsh cinquefoil xyzzzz x x common horsetail xyzzzxyxxxx x few-flowered sedge x xy xyzz x x xy xyzz x bog-rosemary xy xy xy xyzz xyz xyzz crowberry x xy xyz xyzz xyzzz shore sedge xyzzz xy xyzz x x x buckbean xyzz xy xy xyzz xyzz xyzz xyz narrow-leaved cotton-grass xyzz xy xyz xyz xyzz xyz xyz xyzz western bog-laurel xyzzz x scheuchzeria xy xyz x xy great sundew x xy x x xyz xy xyzz xyz round-leaved sundew x x xy xyz xy three-leaved goldthread xyz x xy many-flowered sedge x x xyz xyz deer-cabbage xyz xyz xy pale sedge x xyzz xyzzz xyz great burnet x xy x xyz xyz xy sticky false-asphodel x xyz xyzzzz xyzzzz tufted clubrush xy xy xyzz xy white beak-rush x x xyz xy Alaska bentgrass xyzzz xyzzzz xyzzz xyzzz xyzzzz xyzzzz xyzzzz xyzz xy peat-moss Group I xyzz xyz xy x xy xy x xyz red-stemmed feathermoss xyzz x x x x step moss xyzz xy xyz xy x xy x glow moss xyz xyz xy x golden fuzzy fen moss x xy xy xy xy xy xyzzzz xyzzz xyzz peat-moss Group III xxxx xxyxyzzzxyzreindeer lichens xx xxyzzxyclad lichens xxxyxyzzx peat-moss Group IV xyz xyzzz xyzz hoary rock-moss x xyzz x northern waterfingers xy x bristly swan-neck moss

Chapter 5.1 Bogs 55 Wb01 Black spruce – Creeping-snowberry – Peat-moss

Picea mariana – Gaultheria hispidula – Sphagnum

General Description Black spruce – Creeping-snowberry – Peat-moss bogs are uncommon in the dry and moist SBS and SBPS of the Central and Sub-Boreal Interior at elevations between 500 and 1000 m. Typically, they occur in closed basins or peripheral areas of larger peatlands where there is little groundwater inﬂuence. Picea mariana and Ledum groenlandicum are always present and generally occupy raised microsites. Dwarf woody plants are common and few minerotrophic species are present. Gaultheria hispidula is characteristic and is often prominent. The hummock-forming peat-mosses Sphagnum fuscum and S. capillifolium are dominant in the moss layer but a diversity of other mosses also occurs. On sites with high tree cover, feathermosses can replace Sphagnum as the dominant component of the moss layer. Soils are Mesisols and Fibrisols with a poorly decomposed, acidic, Sphagnum surface tier. Deep sedge and wood peat frequently underlies the surface tier and comprises the bulk of the peat proﬁle.

Characteristic Vegetation Wetland Edatopic Grid

Tree layer (0 - 9 - 50) Soil Nutrient Regime Picea mariana ABCD E F Shrub layer (5 - 35 - 90) Ledum groenlandicum,Picea mariana VM Herb layer (2 - 17 - 50) Gaultheria hispidula,Oxycoccus oxycoccos W Moss layer (70 - 95 - 100) AkVA N MA SA Aulacomnium palustre,Pleurozium schreberi, Sphagnum Group I VW

Soil Moisture Regime pH Comments

St Sl Mo Dy VD The Wb01 represents the “true bog”or “climax” Hydrodynamic Index condition of long-term peatland succession in the sub-boreal forests.A peatland successional sequence in inﬁlling basins appears to follow: Wm01 >> Wf01 >> Wb05 >> Wb01. The related and more common Wb05 occurs where Sphagnum peat accumulation has not yet raised the soil surface well above the groundwater table,and where minerotrophic indicators occur in abundance.Regional climatic conditions likely limit widespread development of Wb01. In small closed basins,Wb01 communities can dominate an entire wetland.However,more commonly,they occur in locations peripheral to Wf02,Wb05,or Wb08. The Wb01 includes only Site Series SBSdk/09 but also occurs elsewhere in the dry/moist SBS and SBPS.

56 Wetlands of British Columbia: A Field Guide to Identiﬁcation Lodgepole pine – Bog rosemary – Peat-moss Wb02

Pinus contorta – Andromeda polifolia – Sphagnum

General Description Lodgepole pine – Bog rosemary – Peat-moss bogs are scattered throughout the wet regions of the Central and Sub- Boreal Interior (ICHvk/wk and SBSvk/wk) at elevations below 1100 m. They occur in closed basins, isolated zones in larger peatlands, and occasionally around acidic peatland lakes. Pinus contorta usually dominates in the sparse and stunted canopy but Picea mariana, Tsuga heterophylla, or Abies lasiocarpa can form a signiﬁcant component. Dwarf woody plants such as Kalmia microphylla and Andromeda polifolia are prominent. On some sites Carex pauciflora and Eriophorum angustifolium may also be prevalent. Soils are Fibrisols and Mesisols typically composed of deep Sphagnum peat but sometimes they occur on peat veneers over dense, ﬁne-textured glaciolacustrine deposits.

Characteristic Vegetation Wetland Edatopic Grid

Tree layer (0 - 5 - 40) Soil Nutrient Regime Picea mariana,Pinus contorta ABCD E F Shrub layer (6 - 32 - 95) Betula nana,Ledum groenlandicum, VM Picea mariana,Pinus contorta Herb layer (3 - 30 - 95) W Andromeda polifolia,Carex pauciﬂora, AkVA N MA SA Eriophorum angustifolium,Kalmia microphylla,Oxycoccus oxycoccos,Rubus VW chamaemorus Moss layer (35 - 95 - 100) Soil Moisture Regime pH Pleurozium schreberi,Sphagnum Group I St Sl Mo Dy VD Hydrodynamic Index Comments The Wb02 Site Association is one of several Bog Site Associations that occur in the interior rainforest.The Wb02 likely reﬂects the “true bog”or “climax”condition of long-term peatland succession in the cool and warm subzones of the interior rainforest of the Central and Sub- Boreal Interior.There are hemlock-dominated bogs (Wb04) in colder and snowier subzones and black spruce–dominated bogs in areas with more boreal climates (such as frost pockets). The Wb07 may be the successional precursor to the Wb02. The Wb02 includes Site Series ICHmm/07,ICHvk2/07,SBSvk/08,and SBSwk3/05 but occurs more widely in the wet SBS and ICH.

Chapter 5.1 Bogs 57 Wb03 Black spruce – Lingonberry – Peat-moss

Picea mariana – Vaccinium vitis-idaea – Sphagnum

General Description Black spruce – Lingonberry – Peat-moss bogs are widespread in the Taiga and Boreal Plains and uncommon in the North- ern Boreal Mountains in topographic depressions with little groundwater inﬂuence. Stunted Picea mariana, usually less than 10 m tall, is always present over an open herb layer and a continuous Sphag- num blanket. Ledum groenlandicum, Rubus chamaemorus, and Vaccinium vitis-idaea are the most abundant understorey species. Sites are hummocky, but because of luxuri- ant Sphagnum growth, hollows are generally no wetter than hummocks and support few minerotrophic indicators. High tree cover on some sites shades out Sphagnum, and feathermosses become dominant. Surface peat on elevated hummocks or domes may dry out and become dominated by Cladonia and Cladina lichens on some sites. Many Wb03 sites are underlain with permafrost and have a domed surface shape. Deep blankets of acidic Sphagnum peat are typical and there is little or no surface water present. Soil types are Fibrisols or Organic Cryosols.

Characteristic Vegetation Wetland Edatopic Grid

Tree layer (0 - 20 - 70) Soil Nutrient Regime Larix laricina,Picea mariana ABCD E F Shrub layer (1 - 35 - 90) Ledum groenlandicum,Picea mariana VM Herb layer (1 - 20 - 99) Equisetum arvense,Rubus chamaemorus, W Vaccinium vitis-idaea AkVA N MA SA Moss layer (48 - 91 - 100) Cladina spp.,Hylocomium splendens, VW Pleurozium schreberi, Sphagnum Group I Soil Moisture Regime pH Comments St Sl Mo Dy VD Hydrodynamic Index The Wb03 represents the climax condition of long-term peatland succession in boreal climates.Climatic conditions in much of this region are favourable to true bog formation and therefore the Wb03 is widespread in suitable terrain.Northern black spruce bogs can occur as simple landscape units in small closed basins, extensive domed bog landscapes,or as zones within larger fen peatlands. Extensive peatlands in the Taiga Plains are primarily the Wb03, with Wb06 occurring along sluggish peatland streams where some water movement is maintained. The Wb03 includes Site Series BWBSdk1/10,BWBSdk2/07,BWBSmw1/08,and BWBSmw2/08.

58 Wetlands of British Columbia: A Field Guide to Identiﬁcation Western hemlock – Cloudberry – Peat-moss Wb04

Tsuga heterophylla – Rubus chamaemorus – Sphagnum

General Description Western hemlock – Cloudberry – Peat-moss bogs are rare in the cold, snowy subzones of the ICH of the Nass Basin, east of the Coast Mountains at elevations below 750 m. These bogs occur in small, closed basins with little or no groundwater inﬂuence. Stunted Tsuga heterophylla is always prominent, but other conifers often occur with low cover. Dwarf woody plant species, especially Rubus chamaemorus and Kalmia microphylla are the dominant component of the open herb layer. Sphagnum growth is strong, elevating most of the soil surface above the watertable; therefore, few minerotrophic species occur. Fibrisols or Mesisols of poorly decomposed Sphagnum peat underlain by deep sedge or woody peat are typical.

Characteristic Vegetation Tree layer (0 - 3 - 11) Tsuga heterophylla Shrub layer (6 - 25 - 60) Ledum groenlandicum,Picea X, Wetland Edatopic Grid Tsuga heterophylla Herb layer (17 - 25 - 90) Soil Nutrient Regime Cornus canadensis,Empetrum nigrum,Erio- ABCD E F phorum angustifolium,Kalmia microphylla, VM Oxycoccus oxycoccos,Rubus chamaemorus Moss layer (90 - 90 - 100) W Pleurozium schreberi,Sphagnum Group I, AM ANAkVA N MA SA Cladina spp. VW Comments Soil Moisture Regime pH The Wb04 represents the “climax”peatland community in the northern interior rainfor- St Sl Mo Dy VD est of the Nass Basin.This area,though low in Hydrodynamic Index elevation,has extremely high snowfall and relatively cool summers.These conditions limit the black spruce or lodgepole pine that would typically occur under similar site conditions.Most Wb04 sites are small in extent.They occur alone or with Wb13 ecosystems. The Wb04 has not been previously described. Further sampling may indicate that the Wb04 also occurs in the wetter subzones of the SBS.

Chapter 5.1 Bogs 59 Wb05 Black spruce – Water sedge – Peat-moss

Picea mariana – Carex aquatilis – Sphagnum

General Description The Black spruce – Water sedge – Peat-moss Bog/Poor Fen Site Association is common throughout the Sub-Boreal and Central Interior (ICH, SBPS, SBS) at elevations below 1300 m. It is found in small closed basins and peripheral areas of larger peatlands where there is a small amount of lateral and groundwater movement and watertable depression. Sites are strongly hummocky with trees and other common bog species rooting on elevated Sphagnum and Tomentypnum nitens mounds, and minerotrophic indicators such as Carex aquatilis, Equisetum spp., and Comarum palustre rooting in the wetter swales. Betula nana is a common and often dominant low shrub in the Wb05. Stunted Picea mariana is normally the predominant tree species but a component of Pinus contorta occurs on some sites. Soils are typically Mesisols of deep (to 4 m) sedge and wood peat. A surface tier of poorly decomposed Sphagnum moss occurs discontinuously, mainly under the raised hummocks.

Characteristic Vegetation Wetland Edatopic Grid

Tree layer (0 - 8 - 55) Soil Nutrient Regime Picea mariana ABCD E F Shrub layer (4 - 40 - 85) Betula nana,Ledum groenlandicum, VM Picea mariana Herb layer (20 - 52 - 98) W Carex aquatilis,Comarum palustre, AkVA N MA SA Gaultheria hispidula,Oxycoccus oxycoccos Moss layer (25 - 90 - 100) VW Aulacomnium palustre,Pleurozium schre- Soil Moisture Regime beri,Sphagnum Group I, Sphagnum pH Group II, Tomentypnum nitens St Sl Mo Dy VD Hydrodynamic Index Comments The Wb05 Site Association is transitional to the Wb01.It has hummock vegetation similar to the Wb01 and Wf01- or Wf02-like vegetation in wetter swales.This suggests that the Wb05 represents the successional state intermediate between sedge fen and “true”bog.A simple peatland successional sequence to the Wb01 is represented the following progression: Wm01 >> Wf01>> Wf02 >>Wb05>>Wb01.The Wb05 is far more abundant than the Wb01 in the sub-boreal,suggesting that regional climatic conditions limit peatland succession.Wb05 communties are often adjacent to the Wf02. The Wb05 includes several Site Series from the ICH,SBPS,and SBS (see Appendix 4).

60 Wetlands of British Columbia: A Field Guide to Identiﬁcation Tamarack – Water sedge – Fen moss Wb06

Larix laricina – Carex aquatilis – Tomentypnum nitens

General Description The Tamarack – Water sedge – Fen moss is a common Bog/ Poor Fen Site Association of the eastern BWBS. It occurs adjacent to domed bogs along peatland streams, water tracks, or groundwater inﬂow seeps. Sites are hummocky, with tamarack and black spruce growing on elevated sites and sedges rooting in the wet hollows. The watertable remains high throughout the growing season. Larix laricina, up to 15 m in height, dominates the canopy, with Picea mariana also present on many sites. A mixed low-shrub understorey dominated by Betula nana can be well developed. Forbs, dwarf shrubs, and smaller sedges root on the elevated hummocks under the tamarack or black spruce trees. Soils are Mesisols of deep sedge and woody peat. Unlike black spruce bogs (Wb03) that occur in the same region, the Wb06 is rarely underlain by permafrost.

Characteristic Vegetation Wetland Edatopic Grid

Tree layer (0 - 0 - 25) Soil Nutrient Regime Larix laricina ABCD E F Shrub layer (10 - 30 - 67) Betula nana, Larix laricina,Ledum groen- VM landicum,Picea mariana,Salix myrtillifolia, S.pedicellaris W Herb layer (20 - 52 - 90) AkVA N MA SA Carex sitchensis,Maianthemum trifolium, Vaccinium vitis-idaea VW Moss layer (60 - 90 - 95) Aulacomnium palustre,Sphagnum Group I, Soil Moisture Regime pH Tomentypnum nitens St Sl Mo Dy VD Hydrodynamic Index Comments The Wb06 occurs in a climatic region that favours the development of classic black spruce bogs (Wb03).The Wb06 occurs only where some watertable flow is maintained.In subdued terrain,this is mainly along sluggish peatland streams and in mountainous terrain at peatland margins adjacent to slopes. The Wb06 is transitional to fens and would be considered a fen under some definitions of wetland classes.However,there are many floristic similarities to true bogs,and the high cover of Sphagnum suggests placement of this unit within the bog class as defined by this guide. The Wb06 includes only Site Series BWBSdk2/08 but is also common throughout the Boreal and Taiga Plains.A Tamarack – Scrub birch – Buckbean Site Series (BWBSmw2/10) has been described for this region (Delong et al.1990),but it is based on limited plots at a single location and likely represents a recently flooded Wb06 site.

Chapter 5.1 Bogs 61 Wb07 Lodgepole pine – Water sedge – Peat-moss

Pinus contorta – Carex aquatilis – Sphagnum

General Description Lodgepole pine – Water sedge – Peat-moss bogs/poor fens are uncommon in the interior rainforest climates at elevations to 1600 m. They most commonly occur in closed basins or in peripheral areas of larger peatlands where there is some groundwater inﬂuence. Pinus contorta, Picea X, and Abies lasiocarpa are all common in the low canopy. Betula nana and Ledum groenlandicum are generally present and often abundant. Abundant Carex aquatilis is characteristic but a diversity of bog-afﬁliated species occurs on hummocks. Sphagnum capillifolium and S. angustifolium form a nearly continuous moss layer. Soils are deep (to 4 m), fibric or mesic peat blankets with a poorly decomposed, acidic, Sphagnum surface tier. Typic Mesisols and Fibrisols are common soil types. Microtopography is often strongly mounded with hummocks of Sphagnum fuscum and S. capillifolium.

Characteristic Vegetation Wetland Edatopic Grid

Tree layer (0 - 5 - 12) Soil Nutrient Regime Picea X,Pinus contorta ABCD E F Shrub layer (15 - 37 - 85) Abies lasiocarpa,Betula nana,Ledum VM groenlandicum,Lonicera involucrata, Picea X,Pinus contorta W Herb layer (46 - 80 - 95) AkVA N MA SA Carex aquatilis,Cornus canadensis, Empetrum nigrum,Equisetum arvense, VW Oxycoccus oxycoccos Soil Moisture Regime Moss layer (40 - 78 - 100) pH Aulacomnium palustre,Pleurozium schre- St Sl Mo Dy VD beri,Sphagnum Group I,Tomentypnum Hydrodynamic Index nitens

Comments The Wb07 is the southern and wet-climate equivalent of the Wb05 that occurs widely in the SBS and BWBS.The distrbution of Wb07 sites coincides with regions where Picea mariana does not occur.

62 Wetlands of British Columbia: A Field Guide to Identiﬁcation Black spruce – Soft-leaved sedge – Peat-moss Wb08

Picea mariana – Carex disperma – Sphagnum

General Description Spruce – Soft-leaved sedge – Peat-moss bogs/poor swamps are uncommon throughout the Interior below 1700 m in palustrine depressions fed by slow-moving groundwater. These sites are often strongly hummocky; trees and upland species occur on mounds. Stand- ing water is often present between hummocks, but sites are not fully ﬂooded. The coniferous canopy of spruce and black spruce is open. The trees grow poorly because of saturated soils, though there may be large individual stems on some sites. Carex disperma and Equisetum spp. co-dominate on many sites. The moss layer is diverse; peat-mosses, leafy mosses, and feathermosses can all be prominent. On wetter sites, with deep standing water in depressions, Equisetum fluviatile or E. palustre can be abundant. Soils are usually deep (1–4 m) Typic Mesisols or Humisols derived from woody peat but are occasionally thin organic veneers over limnic deposits or ﬁne-textured lacustrine materials.

Characteristic Vegetation Wetland Edatopic Grid

Tree layer (0 - 25 - 55) Soil Nutrient Regime Picea mariana,Picea X ABCD E F Shrub layer (6 - 30 - 80) Betula nana,Ledum groenlandicum, VM Picea mariana,Picea X Herb layer (10 - 64 - 99) W Carex aquatilis,C. disperma,C.tenuiﬂora, AkVA N MA SA Comarum palustre,Equisetum arvense, E. ﬂuviatile VW Moss layer (2 - 76 - 100) Soil Moisture Regime Aulacomnium palustre,Hylocomium pH splendens,Mnium spp.,Pleurozium schreberi, St Sl Mo Dy VD Sphagnum Group I, Tomentypnum nitens Hydrodynamic Index

Comments The Wb08 occupies sites intermediate between the Wb05 and the more productive Ws07. These sites are transitional bogs to swamps,with bog communities occurring on elevated mounds and more minerotrophic species occurring in wet swales.The closely related Wb09 has a more northerly distribution,less standing water,and lower productivity.These ecosystems often occur in peripheral areas of larger peatlands or wet depressions adjacent to shrub swamps. The Wb08 includes many Site Series from the SBS (see Appendix 4) but is now recognized as being more widespread.

Chapter 5.1 Bogs 63 Wb09 Black spruce – Common horsetail – Peat-moss

Picea mariana – Equisetum arvense – Sphagnum

General Description The Black spruce – Common horsetail – Peat-moss Bog/Poor Swamp Site Association is uncommon in the Central Interior and Northern Boreal Mountains (BWBS, SBS) in small palustrine basins and at the periphery of larger peatlands. This Site Association is transitional to forested swamps but has, in contrast, abundant bog-affiliated species, very poor tree growth, and more stagnant hydrology. Sites are often strongly hummocky, with conifers and typical bog species occurring on elevated sites and minerotrophic indicators in hollows. Hummock species include stunted Picea mariana, Ledum groenlandicum, and Sphagnum spp. Equisetum arvense is always present between hummocks. Soils can be deep Sphagnum peat (to 3 m) or shallow veneers over fine-textured mineral materials. Mesisols and Gleysols are equally common. Standing water can persist between hummocks, but elevated sites are never flooded.

Characteristic Vegetation Wetland Edatopic Grid

Tree layer (0 - 17 - 77) Soil Nutrient Regime Picea mariana ABCD E F Shrub layer (10 - 37 - 95) Betula nana, Ledum groenlandicum, VM Picea mariana Herb layer (18 - 58 - 95) W Calamagrostis canadensis,Cornus canaden- AkVA N MA SA sis,Equisetum arvense Moss layer (18 - 58 - 95) VW Aulacomnium palustre,Hylocomnium palustre,Mnium spp.,Pleurozium schreberi, Soil Moisture Regime pH Sphagnum Group I,Tomentypnum nitens St Sl Mo Dy VD Hydrodynamic Index Comments The Wb09 is the northern equivalent of the Wb08 and represents sites transitional between the Wb03 and Ws07. The Wb09 includes Site Series BWBSdk1/09, wk1/07,and wk2/07 also occurs elsewhere in the BWBS and northern SBS.

64 Wetlands of British Columbia: A Field Guide to Identiﬁcation Lodgepole pine – Few-ﬂowered sedge – Peat-moss Wb10

Pinus contorta – Carex pauciﬂora – Sphagnum

General Description Lodgepole pine – Few-flowered sedge – Peat-moss bogs/poor fens are rare at montane elevations in the Sub-Boreal Interior and Southern Interior Mountains. These ecosystems occur as small stands in frost-prone basins or on gradual slopes. Pinus contorta is always present as a sparse canopy. Trees are small but well formed and are not rooted on elevated microsites. The shrub layer consists almost entirely of stunted conifers, giving these sites an open, park-like character. Carex pauciflora usually dominates the herb layer but there is a diversity of other graminoids and typical bog dwarf shrubs. The moss layer is most often a continuous lawn of Sphagnum angustifolium with scattered other species. This Site Association usually has a smooth microtopography and is saturated at the surface from seepage. Soil water is moderately acid, suggesting that groundwater inputs are poor in minerals. Soils are Typic Humisols and Mesisols with a surface tier of poorly decomposed Sphagnum peat.

Characteristic Vegetation Wetland Edatopic Grid

Tree layer (0 - 1 - 22) Soil Nutrient Regime Pinus contorta ABCD E F Shrub layer (3 - 20 - 45) Betula nana,Ledum groenlandicum, VM Pinus contorta Herb layer (15 - 44 - 75) W Carex aquatilis,C. pauciﬂora,Eriophorum AkVA N MA SA angustifolium,Kalmia microphylla, Oxycoccus oxycoccos VW Moss layer (30 - 85 - 100) Soil Moisture Regime Pleurozium schreberi, Sphagnum Group I pH

St Sl Mo Dy VD Comments Hydrodynamic Index Few Wb10 sites have been sampled; the site conditions that support this Site Association are not well understood and appear to be uncommon.A high watertable and limited microtopography would suggest limited potential for tree growth but many of these sites have trees to 15 m. This ecosystem has been found most often as a pure stand type not associated with other wetland ecosystems,but may border Wf01 or Wf12 sites.

Chapter 5.1 Bogs 65 Wb11 Black spruce – Buckbean – Peat-moss

Picea mariana – Menyanthes trifoliata – Sphagnum

General Description The Black spruce – Buckbean – Peat-moss Bog Site Associa- tion is uncommon in the wet climates of the Sub-Boreal Interior and the Nass Basin at elevations below 1200 m. These sites are found in small inﬁlled basins or on edges of larger peatlands where the watertable is stagnant. Picea mariana and/or Pinus contorta are always present. Tree cover and growth are very sparse and stunted (< 2 m) on wetter sites but cover and growth increase with declining watertable. A diverse assemblage of graminoids and shrubs is typical, with Menyanthes trifoliata always prominent. Sites can be hummocky with some standing water in depressions, or with a dense, continuous Sphagnum lawn. Soils are commonly deep moss- and sedge-derived peat. Typic Mesisols and Fibrisols are the most common soil types but Terric subgroups or Humisols derived from limnic materials also occur.

Characteristic Vegetation Wetland Edatopic Grid Tree layer (0 - 8 - 60) Picea mariana,Pinus contorta Soil Nutrient Regime Shrub layer (5 - 28 - 85) ABCD E F Betula nana,Ledum groenlandicum, VM Picea mariana,Pinus contorta Herb layer (8 - 53 - 90) W Carex aquatilis,C.limosa,Comarum palustre,Equisetum ﬂuviatile,Menyanthes AkVA N MA SA trifoliata,Oxycoccus oxycoccos VW Moss layer (20 - 95 - 100) Aulacomnium palustre,Pleurozium Soil Moisture Regime pH schreberi,Sphagnum Group I, Sphagnum Group II,Tomentypnum nitens St Sl Mo Dy VD Hydrodynamic Index Comments Wb11 develops from Wf07 or Wb13 ecosystems where Menyanthes trifoliata is a prominent species.It is likely that M. trifoliata is a persistent species that initially establishes during the early phases of basin-inﬁlling and contin- ues to grow apace of peat accumulation.

66 Wetlands of British Columbia: A Field Guide to Identiﬁcation Scheuchzeria – Peat-moss Wb12

Scheuchzeria palustris – Sphagnum

General Description Scheuchzeria – Peat-moss bogs are uncommon in the sub- boreal and boreal forests at elevations below 1000 m. They usually occur as small inclusions in larger peatlands on floating mats with continually saturated peat and restricted water movements. Vegetation is characterized by species tolerant of permanent saturation but intolerant of deep flooding. A low shrub layer of Salix pedicellaris occurs on some sites but dwarf shrubs such as Andromeda polifolia, Kalmia microphylla, and Oxycoccus oxycoccos are more prevalent. Scheuchzeria palustris is always prominent and Carex limosa occurs on most sites. The moss layer is dominated by Sphagnum Group I species. Soils are mostly fibric Sphagnum peat and can be floating mats. The watertable is at the surface but does not flood more than several centimetres above the soils surface. The water is very stagnant and low in dissolved oxygen.

Characteristic Vegetation Wetland Edatopic Grid

Tree layer (0 - 0 - 0) Soil Nutrient Regime Shrub layer (0 - 8 - 20) ABCD E F Salix pedicellaris Herb layer (20 - 37 - 90) VM Andromeda polifolia,Carex limosa, Eriophorum chamissonis,Kalmia W microphylla,Oxycoccus oxycoccos, AkVA N MA SA Scheuchzeria palustris. VW Moss layer (60 - 67 - 100) Soil Moisture Regime Sphagnum Group I pH

St Sl Mo Dy VD Comments Hydrodynamic Index The Wb12 Site Association requires permanent saturation with acidic waters in combination with no ﬂooding.It therefore occurs in wetlands where water regimes are relatively stable and on sites with ungrounded peat that can rise and fall with changes in watertable.On sites with higher pH,the Wb12 is replaced by the Wf07 or Wf08. In the eastern boreal areas,ecosystems that are very similar to Wb12 but have Sarracenia purpurea and Chamaedaphne calyculata occur (see additional units).

Chapter 5.1 Bogs 67 Wb13 Shore sedge – Buckbean – Peat-moss

Carex limosa – Menyanthes trifoliata – Sphagnum

General Description Shore sedge – Buckbean – Peat-moss bogs are uncommon in the interior rainforest and coastal transition regions at elevations below 1600 m. They occur as components of larger acidic peatlands, occupying the central, wettest portions of the peatland: either grounded, highly saturated peat blankets, or ﬂoating mats. Species tolerant of acidic, continually saturated conditions and concurrent lack of oxygen are prominent. The most consistent of these is Carex limosa. Drosera anglica, Menyanthes trifoliata, Kalmia microphylla, and other species can be abundant, sparse, or absent on Wb13 sites. Sphagnum angustifolium, S. magellanicum, or S. fuscum often form a continuous lawn or there may be a mix of species in hummock-hollow patterns. Soils are deep (to > 5 m) sedge-derived Mesisols with a surface tier of poorly decomposed Sphagnum peat. The watertable is typically at or near the surface but there is little standing water.

Characteristic Vegetation Wetland Edatopic Grid Tree layer (0 - 0 - 0) Shrub layer (0 - .5 - 10) Soil Nutrient Regime Herb layer (15 - 73 - 100) ABCD E F Carex limosa,Drosera anglica,Eriophorum VM angustifolium,Kalmia microphylla, Menyanthes trifoliata,Trientalis europaea W ssp. arctica Moss layer (30 - 90 - 100) AkVA N MA SA Sphagnum Group I VW

Comments Soil Moisture Regime pH

The Wf08 is a similar unit that occurs in St Sl Mo Dy VD drier interior climates on saturated sites. Hydrodynamic Index This unit most frequently occurs in wetter locations adjacent to the Wb11 or Wb02, and beside peatland ponds or lakes.

68 Wetlands of British Columbia: A Field Guide to Identiﬁcation Labrador tea – Bog-laurel – Peat-moss Wb50

Ledum groenlandicum – Kalmia microphylla – Sphagnum

General Description Labrador tea – Bog laurel – Peat-moss bogs occur uncom- monly in the drier subzones of the south Coast at low to montane elevations. They are raised bogs in closed basins with a high, stagnant watertable or adjacent to peatland lakes. Some locations may be on ﬂoating mats. The vegetation is low in stature and dominated by Ledum groenlandicum with an abundance of Kalmia microphylla and Oxycoccus oxycoccos. Myrica gale or dwarfed Pinus contorta var. contorta can be prominent on some, usually drier, sites. Herb cover is variable, low-lying areas can have a high cover of Rhynchospora alba while raised sites can have Rubus chamaemorus in abundance. Group I Sphag- num spp. are most common (S. fuscum, S. capillifolium) but coastal species also occur (S. papillosum), mostly in wetter hollows. Soils are Typic Fibrisols or Mesisols with surface tier of poorly decomposed Sphagnum peat.

Characteristic Vegetation Wetland Edatopic Grid Tree layer (0 - 0 - 0) Shrub layer (0 - 20 - 85) Soil Nutrient Regime Ledum groenlandicum,Myrica gale, ABCD E F Pinus contorta VM Herb layer (10 - 30 - 50) Drosera rotundifolia,Kalmia microphylla, W Oxycoccus oxycoccos,Rhynchospora alba Moss layer (50 - 90 - 91) AkVA N MA SA Sphagnum Group I VW

Comments Soil Moisture Regime pH

The Wb50 is widespread but generally of St Sl Mo Dy VD small extent except for several notably ex- Hydrodynamic Index tensive bogs in subdued terrain (e.g.,Burns Bog).It occurs in open,unshaded locations adjacent to other low-stature peatland types or open water. The microtopography of this unit is often broken by peat degradation hollows caused by die- back of Sphagnum and increased localized decomposition.Shallow pools caused by peat degradation are frequent and are occupied by species such as Menyanthes trifoliata, Scheuchzeria palustris,or Nuphar lutea.

Chapter 5.1 Bogs 69 Wb51 Shore pine – Black crowberry – Tough peat-moss

Pinus contorta var. contorta – Empetrum nigrum – Sphagnum austinii

General Description Shore pine – Black crowberry – Tough peat-moss are raised bogs that occur on level terrain or topographic depressions in the Coast and Mountains at elevations below 100 m. “Bonsai” Pinus contorta are scattered with other low shrubs such as Chamaecyparis nootkatensis, Myrica gale, and Juniperus communis. Empetrum nigrum, Kalmia microphylla, and Rubus chamaemorus are typical prominent dwarf shrubs. Several Sphagnum species occur in these bogs but the distinctive Sphag- num austinii, which forms dense tough mounds, is very common, especially in northern sites. Soils are commonly Mesisols of Sphagnum peat, fibric at the surface and mesic or humic at depth. Sites are slightly domed and raised above surrounding sites through the active growth of Sphagnum. Peat depths range from 0.5 m to > 4 m.

Characteristic Vegetation Wetland Edatopic Grid

Tree layer (0 - 0 - 5) Soil Nutrient Regime Shrub layer (7 - 30 - 65) ABCD E F Chamaecyparis nootkatensis,Juniperus communis,Ledum groenlandicum,Myrica VM gale,Pinus contorta,Thuja plicata Herb layer (13 - 35 - 50) W Carex livida,C.pluriﬂora,Empetrum AkVA N MA SA nigrum,Eriophorum angustifolium, Kalmia microphylla,Rubus chamaemorus, VW Sanguisorba ofﬁcinalis,Triantha glutinosa, Trichophorum cespitosum Soil Moisture Regime pH Moss layer (90 - 95 - 100) St Sl Mo Dy VD Racomitrium lanuginosum, Hydrodynamic Index Sphagnum Group I,Sphagnum Group III

Comments The Wb51 represents actively growing bogs that show few signs of the stagnation in peat accumulation (such as high abundance of ground lichens and peat degradation pools) common in other bogs of the outer Coast.Extensive areas of the Wb51 can be found in and around Naikoon Provincial Park on the Queen Charlotte Islands where large tracts of domed bog have developed over marine or glacial outwash sediments.Elsewhere,this community type occurs as smaller areas in the blanket mire complex or in small topogenous bogs. The Wb51 includes Site Series CWHvh2/31 and CWHwh1/11.

70 Wetlands of British Columbia: A Field Guide to Identiﬁcation Common juniper – Tufted clubrush – Hoary rock-moss Wb52

Juniper communis – Trichophorum cespitosum – Racomitrium lanuginosum

General Description Common juniper – Tufted clubrush – Hoary rock-moss bogs are a very common component of the “blanket mire complex” of the outer Coast at elevations below 800 m. Scattered, “bonsai” shore pine are always present but the shrub layer is characterized more by Juniperus communis and Myrica gale. Tri- chophorum cespitosum is always dominant but there is a diversity of dwarf shrubs and herbs. On sites with deeper peat, Sphagnum spp. are co-dominant with Racomitrium lanuginosum, Cladina spp., and Siphula ceratites. On sites with shallow peat, moss layer cover is lower, Sphagnum is greatly reduced, and Siphula ceratites and Campylopus atrovirens become more prominent. Deposits of dark mesic peat to 1.5 m, underlain by bedrock, are typical, but peat depth varies considerably. Some sites on the extreme outer Coast have a complex of poorly drained mineral soils derived from bedrock and discontinuous organic accumulations 5–50 cm deep.

Characteristic Vegetation Wetland Edatopic Grid

Shrub layer (0 - 27 - 90) Soil Nutrient Regime Chamaecyparis nootkatensis,Juniper com- ABCD E F munis,Ledum groenlandicum,Myrica gale, Pinus contorta,Vaccinium uliginosum VM Herb layer (8 - 70 - 99) Agrostis aequivalvis,Andromeda polifolia, W Carex livida,Coptis trifolia,Drosera rotundifo- AkVA N MA SA lia,Empetrum nigrum,Eriophorum angustifolium,Fauria crista-galli,Kalmia VW microphylla,Rhynchospora alba,Sanguisor- Soil Moisture Regime ba ofﬁcinalis,Trichophorum cespitosum pH Moss layer (5 - 75 - 99) St Sl Mo Dy VD Campylopus atrovirens,Cladina spp., Hydrodynamic Index Racomitrium lanuginosum,Siphula ceratites,Sphagnum Group I,Sphagnum Group III,Sphagnum Group IV

Comments The Wb52 probably represents “over-mature”bogs where Sphagnum mosses are limited and peat is no longer accumulating.This is reﬂected in the prevalence of Racomitrium lanuginosum and ground lichens,which would not normally be able to compete with Sphagnum. Furthermore,on most sites there are circular to teardrop-shaped shallow pools that are created by peat degradation.These pools are 1–4 m2 in area and 10–50 cm deep,ringed by dams of Sphagnum and Trichophorum cespitosum growing in strongly tenacious peat. The Wb52 includes Site Series CWHvh2/32.

Chapter 5.1 Bogs 71 Wb53 Shore pine – Yellow-cedar – Tufted clubrush

Pinus contorta – Chamaecyparis nootkatensis – Trichophorum cespitosum

General Description The Shore pine – Yellow-cedar – Tufted clubrush Site Associ- ation is a very common component of the “blanket mire complex” of the outer Coast. The Wb53 occurs on gently to steeply sloping terrain on slightly shedding sites such as hillocks and slope breaks, or simply along drainageways. There is rarely standing water on these sites. Stunted yellow-cedar and shore pine to 10 m is characteristic and differentiates this association from the Wb52. Trichophorum cespitosum is prominent in the understorey but there is a diversity of other herbs and dwarf shrubs. The moss layer is moderately well developed with a mix of upland species on elevated sites and Sphagnum spp. in wetter hollows. Soils are variable, ranging from deep fibric sedge and wood peat deposits (> 2 m) to thin peat veneers over granitic bedrock. Hummocky microtopography provides drier sites for tree establishment. Wetland Edatopic Grid Characteristic Vegetation Soil Nutrient Regime Tree layer (0 - 0 - 10) ABCD E F Pinus contorta Shrub layer (10 - 55 - 95) VM Chamaecyparis nootkatensis,Gaultheria shallon,Ledum groenlandicum,Pinus con- W torta,Thuja plicata AkVA N MA SA Herb layer (25 - 70 - 95) Cornus canadensis,Drosera rotundifolia, VW

Empetrum nigrum,Eriophorum angustifoli- Soil Moisture Regime pH um,Fauria crista-galli,Kalmia microphylla, Sanguisorba ofﬁcinalis,Trichophorum ce- St Sl Mo Dy VD spitosum,Vaccinium uliginosum Hydrodynamic Index Moss layer (5 - 50 - 90) Cladina spp.,Pleurozium schreberi, Racomitrium lanuginosum,Sphagnum Group I,Sphagnum Group III

Comments The Wb53 has been refered to as “bog woodland”and is a major component of the blanket mire complex of the outer Coast.It typically occurs with the related Wb52 on sloping terrain. The Wb53 includes Site Series CWHvh2/12.

72 Wetlands of British Columbia: A Field Guide to Identiﬁcation Additional Bog Site Associations

This section briefly describes some uncommon Bog Site Associations that have been sampled in British Columbia. Western redcedar – White pine – Bristle-stalked sedge Thuja plicata – Pinus monticola – Carex leptalea Western redcedar – White pine – Bristle-stalked sedge stands are uncommon in the Georgia Depression in regions underlain with basic bedrock. They occur in small depressions and around peatland lakes on raised peat deposits. A diverse mixture of conifers forms an open, tall-shrub layer. Western redcedar and white pine are common but western hemlock, Sitka spruce, and shore pine also occur. Labrador tea and sweet gale are common in the shrub layer. Bristle-stalked sedge is always prominent in the herb layer but other sedges can be common. Peat-mosses predominate; sites with bog-like vegetation but with brown mosses instead of peat-mosses have been observed. Leatherleaf – Pitcher plant – Peat-moss Chamaedaphne calyculata – Sarracenia purpurea – Sphagnum Leatherleaf – Pitcher plant – Peat-moss sites have been observed but infrequently sampled in the Taiga Plains Ecoprovince. They occur on floating mats or other peatland locations where the watertable is maintained at, but not above, the peat surface. These communities are very similar in species composition and structure to the Wb12 of the western boreal and sub-boreal regions. However, leatherleaf (Chamaedaphne calyculata) replaces bog willow (Salix pedicellaris), and pitcher plant (Sarracenia purpurea) occurs. The Taiga Plains Ecoprovince represents the western limit of pitcher plant in the Canadian boreal forest. Shore pine – Labrador tea – Salal Pinus contorta var. contorta – Ledum groenlandicum – Gaultheria shallon Shore pine – Labrador tea – Salal stands are uncommon in the Georgia Depression in small topographic depressions and around the edge of domed bogs. Shore pine is often the only tree species in an open and stunted (<10 m) canopy. Ledum groenlandicum and Gaultheria shallon typically form a dense, low-shrub layer. Kalmia polifolia, Oxycoccus oxycoccos, Pteridium aquilinium, and Cornus canadensis are common in the typically sparse

Chapter 5.1 Bogs 73 Additional Bog Site Associations

herb layer. Sphagnum capillifolium is often a small component of the moss layer, with feathermosses, such as Hylocomium splendens, dominating on most sites. Soils have a surface horizon of feathermoss and litter-derived humus over deep peat deposits composed of Sphagnum remains.

1 Sphagnum spp., peat moss 2 Ledum groenlandicum, Labrador tea 3 Oxycoccus oxycoccos, bog cranberry

74 Wetlands of British Columbia: A Field Guide to Identiﬁcation 5.2 FENS

1 A patterned fen near Williston Reservoir, northern Rocky Mountains (SBSmk2) 2 A basin fen near Babine Lake, Sub-Boreal Interior (SBSmc2) 3 Sloping fens, Harold Price Plateau near Smithers (ESSFwv)

Chapter 5.2 Fens 75 FEN WETLAND CLASS

Definition A fen is a nutrient-medium peatland ecosystem dominated by sedges and brown mosses, where mineral-bearing groundwater is within the rooting zone and minerotrophic plant species are common. General Description Vegetation Table 5.2.2 lists species common to Fen Site Associations described in this guide. Fens are characterized by high cover of sedges and bryophytes of the brown moss group, such as Campylium, Drepanocladus, Scorpidi- um, Tomentypnum, and Warnstorfia. These species reflect relatively mineral-rich site conditions in fens compared to bogs. Some sites may have high cover of minerotrophic non–hummock-forming Sphagnum species (see Group II Sphagnum definition page 47). Low shrub or graminoid physiognomy is typical: a high watertable precludes tall shrubs and trees. Shrub species are typically deciduous, most commonly willows and scrub birch. On very saturated sites some evergreen dwarf shrubs such as bog cranberry and bog rosemary may be prominent. The moss layer is usually well developed. Landscape Position and Distribution Fens develop where permanently saturated soil conditions are maintained. Common locations for fens are groundwater-fed basins, gradual seepage slopes, and protected lake or pond margins where there is little wave action or drawdown. Fens are the most common wetland class in the province and occur in all but the warmest and driest climates (Table 5.2.1). Regions with cool summer temperatures are optimal for fen formation.

76 Wetlands of British Columbia: A Field Guide to Identification Hydrology and Soils Fens occur where peat accumu- Soil Nutrient Regime lates but where groundwater ABCD E F flows maintain relatively high M mineral content in the rooting VM zone. The watertable is usually at or near the soil surface with little W late season drawdown. The degree AkVA N MA SA of lateral flow, stability of the wa- VW Fens tertable, and availability of base Soil Moisture Regime cations differentiate most fen pH ecosystems (Figure 5.2.1). St Sl Mo Dy VD Hydrodynamic Index Soils are of the Organic order,  .. most frequently Mesisols, derived Position of fens on the edatopic grid. from sedges and mosses. Soil nutrient regime ranges from poor to rich. Other Comments In long-term peatland development, fens are classically represented as an intermediate stage between marsh (with mineral rooting substrate) and bog ecosystems (semi-terrestrial peatlands). The peat profile and vegetation of some peatland ecosystems in British Columbia do reflect this model; the lowest tier of peat indicates marsh conditions, the middle tier is sedge-dominated, and the upper tier is Sphagnum-derived. However, many fens occur in climatic areas that are not conducive to bog development and therefore may be long-lived. In addition, some fens have peat profiles with consistent peat composition throughout, suggesting that they have been stable ecosystems since peatland initiation and are not undergoing long-term peatland succession. Conservation Issues Fens are the most common wetland class throughout most of British Columbia. Site Associations dominated by water, beaked, or Sitka sedges and narrow-leaved cotton-grass (Wf01–03, 11, 12) are the most common of the fen types in the province in part because these species tolerate a wide range of hydrologic conditions. These large sedge–dominated Site Associations persist despite beaver flooding, beaver dam removal, hay- ing, extreme flooding, sedimentation, or burning. However, fundamental changes to the water regime such as permanent watertable elevation or draining will convert communities to other types.

Chapter 5.2 Fens 77 Fen Site Associations with high stable water regimes (Wf05 – Wf11) are less common and more sensitive to lowered watertables. They are re- silient to moderate increases in watertable since peat blankets will often swell or float in response, thereby preventing deep flooding. Fens have moderate wildlife habitat values. Forage availability is moderate for ungulates and very low for bear in most fens. Fens with standing water will support moderate aquatic insect populations, which in turn support avian and mammalian insectivores. Shrew and some rodents such as jumping mice and voles will use fens extensively. Fens with relatively stable water regimes and well-developed bryophyte layers support unique peatland arthropod communities. Waterfowl use of fens is often limited because of low prey abundance. Other waterbirds such as Common Snipe and Black Tern use fens for nesting. Agricultural uses of fens are primarily for grazing and the production of “swamp hay.” The Wf01 is the most commonly used Site Association for this purpose. Wetter fens are generally of low value because of trafficabil- ity problems associated with saturated peat soils.

Drepanocladus aduncus, common hook-moss

78 Wetlands of British Columbia: A Field Guide to Identiﬁcation s s i x xxx xxxxx xx BGPP BWBS SWB ESSF ICH IDF MS SBS CDF CWH SBPS MH Distribution of Fen Site Associations by biogeoclimatic zone Water sedge – Beaked sedge sedge – Beaked Water sedgeWater – Scrub birch sedge – Peat-mossWater mosss sedge – Glow Water willow – Barclay’s hook-mossSlender sedge – Common Slender sedge – Buckbean sedge – Shore – Buckbean Scrub birch – Hook-moss sedge – Buckbean Shore – Hook-moss spike-rush Few-ﬂowered hook-moss clubrush – Red Bay Hudson clubrush – Star mossTufted – Marsh-marigold cotton-grass Narrow-leaved sedge – Shore cotton-grass Narrow-leaved – Peat-moss cotton-grass Narrow-leaved x sedge – Peat-mossSitka sedge gale – Sitka Sweet xxx beak-rush White Slender sedge – xx x x xxx x x x xx xx xx xxx xx x x x xxx xx xx x xxx x x xxx xx xx x xx x xx xx x x x x x x x x x x x x x x x x x x xx xx i = inland areas only s = southern subzones only x = incidental; < 5% of wetlands xx = minor; 5–25% of wetlands xxx = major; >25% of wetlands Wf01 Wf02 Wf03 Wf04 Wf05 Wf06 Wf07 Wf08 Wf09 Wf10 Wf11 Wf12 Wf13 Wf50 Wf51 Wf52 Wf53  .. 

Chapter 5.2 Fens 79  .. Fen Species Importance Table

Species Wf01 Wf02 Wf03 Wf04 Wf05 Wf06 Wf07 Wf08 Shrubs Betula nana x xyzzz x xy x x xyzzz x Salix barclayi x x xyzzzz Salix pedicellaris x xyz xy xy xyzz xy Spiraea douglasii xxxx Myrica gale Herbs Carex utriculata xyzzz xyz xy xy xyz x x x and Carex aquatilis xyzzz xyzz xyzzz xyzz xyz x xyz x Dwarf Comarum palustre xy xyz xy xy xy xyzz xy Shrubs Calamagrostis canadensis x xy x xyzz x x x x Carex lasiocarpa x xyzzzz xyzzzz xy x Menyanthes trifoliata x x x xyzzz xyzzz xyzz Carex limosa x x x x xy xyzz xyzzz Carex chordorrhiza x x xy xyz xy Eleocharis quinqueflora xx Trichophorum alpinum x Trichophorum cespitosum Eriophorum angustifolium xxyxyxxxyxy Caltha leptosepala xy xy Carex anthoxanthea Equisetum fluviatile x x xy xy xy xy Carex magellanica xx Carex sitchensis xy xy xyz xyz x x Rhynchospora alba x Carex livida xxxx Eriophorum chamissonis xxx Vahlodea atropurpurea x Drosera anglica xxxy Hypericum anagalloides Triantha glutinosa xxxy Schoenoplectus tabernaemontani Fauria crista-galli Senecio triangularis x x xyz xy Andromeda polifolia xxyxy Kalmia microphylla xxx Oxycoccus oxycoccos xx xxxyx Triglochin maritima xxxxyxy Drosera rotundifolia xx x Leptarrhena pyrolifolia xy x Platanthera dilatata xxy xxx Sanguisorba canadensis xxy Utricularia intermedia xx xxy Viola palustris xx x Lichens Sphagnum Group I xy xyz xyzz x x xyz x and Aulacomnium palustre x xyz xyz xyzz x x x x Mosses Drepanocladus spp. xyz x xy xyzz xyz xyz xyz Sphagnum Group II x xyz x x xyz xy Tomentypnum nitens x xyzz xyz xy x xyz xy Philonotis fontana xy xyz Calliergon stramineum xxxxx Scorpidium spp. x xy xyzz xy Campylium stellatum x x xy xy x x Warnstorfia spp. x xy x x xyz x xy Meesia triquetra x x xy xy

80 Wetlands of British Columbia: A Field Guide to Identification Wf09 Wf10 Wf11 Wf12 Wf13 Wf50 Wf51 Wf52 Wf53 Common Name xxxyx scrub birch xx Barclay’s willow xy x x bog willow x xyz x pink spirea x xyzzzz xyz sweet gale xyxxxyx x beaked sedge x xyz x xy xyz x x water sedge xy x x x xyz xy x marsh cinquefoil x x xy xy xy xy x bluejoint reedgrass x xyz xy x x x xyzzz slender sedge xyzzzxyzxxxxyxx buckbean xyz xyzz xyz x xyzz x x shore sedge xyzz x cordroot sedge xyzzzz x x few-flowered spike-rush xyzzzz x Hudson Bay clubrush x xyzzzz x xyz x tufted clubrush xyz x xyz xyzzzz xyzzz xyzzzz xy x narrow-leaved cotton-grass xy x xyzz xyz x x white mtn. marsh-marigold xy x yellow-flowered sedge xxx swamp horsetail x x xyz x poor sedge xxxxyxxyzzzz xyzzz x Sitka sedge xy xyzz white beak-rush xy x x pale sedge xxyChamisso’s cotton-grass xy xy x mountain hairgrass x xyzz xy x x x xy great sundew xx bog St. John’s-wort x xyz xy xy xy sticky asphodel xy great bulrush xyz x deer-cabbage x xxx x arrow-leaved groundsel xyz x x bog-rosemary x x xy x xyz x x western bog-laurel x xyz x xy x bog cranberry xyz x seaside arrow-grass xxyxxyround-leaved sundew xy xy x x leatherleaf saxifrage xyxxxyxxx fragrant white rein orchid x x x xyxyx Sitka burnet xxy x flat-leaved bladderwort xxxxxyx marsh violet x xy xy x xyz xyzz xyzz x peat-moss Group I xy x x xyz xyz xy xy x glow moss xyz xy xy x x hook-mosses xy xyz xyz x x xyzz x peat-moss Group II xyz xyz x xy x x golden fuzzy fen moss xy xyz spring moss xy xyz x xy x x x straw spear-moss xy xyzz x x x sausage-moss xyzzz xyzzz xy yellow star-moss xxxyxy x hook-mosses xy x x three-ranked hump-moss

Chapter 5.2 Fens 81 Wf01 Water sedge – Beaked sedge

Carex aquatilis – Carex utriculata

General Description The Water sedge – Beaked sedge Fen Site Association is the most common and widespread Fen Site Association in the province. It occurs in all but the warmest and driest subzones from low to subalpine elevations on sites that are annually inundated by shallow, low-energy ﬂood waters and that experience some late-season drawdown. Wf01 fens are found in a wide variety of landscape positions but most commonly palustrine basins.They occupy wetter zones in larger peatland complexes but also form extensive pure “meadows.” Species diversity is low; Carex aquatilis and Carex utriculata cover is often continuous, with scattered forbs, aquatics, and mosses in the understorey. On sites that dry out at the surface, Calamagrostis canadensis or C. stricta can become prominent, species diversity increases, and sites become more meadow-like. Peat depths range from 30 to > 300 cm. Common soil types include typic and terric Fibrisols and Mesisols. This Site Association tolerates variable hydrology. Characteristic Vegetation Wetland Edatopic Grid

Tree layer (0 - 0 - 0) Soil Nutrient Regime Shrub layer (0 - 0 - 10) ABCD E F Herb layer (13 - 80 - 100) Carex aquatilis,C. utriculata VM Moss layer (0 - 5 - 100) Drepanocladus aduncus W AM ANAkVA N MA SA Comments Sites dominated by C. utriculata and C. VW

aquatilis but with mineral or humic soils are Soil Moisture Regime pH described by the Wm01. Because Wf01 and

Wm01 sites are species-poor and the two dom- St Sl Mo Dy VD inant sedge species have a wide ecological Hydrodynamic Index amplitude,the plant community poorly differentiates between sites on peat (Wf01) and those on mineral soil (Wm01).Wf01 sites typically have less C. utriculata and fewer aquatics than Wm01 sites.The Wf01 develops from the Wm01 in most circumstances. Sites that are drier or at least have more pronounced microtopography than the Wf01 are usually occupied by communities with low shrubs and high moss cover (most commonly,the Wf02).However,at higher elevations few shrubs occur and only moss cover increases (Wf03). Sites with greater waterﬂow are characterized by tall-shrub swamps dominated by willows or alders,and water sedges,and have mineral or humic peat soils.

82 Wetlands of British Columbia: A Field Guide to Identiﬁcation Scrub birch – Water sedge Wf02

Betula nana – Carex aquatilis

General Description The Scrub birch – Water sedge Fen Site Association is one of the most common peatland Site Associations throughout the Interior and is absent only from PP/BG and wet ESSF subzones. It is frequently a major component of large peatlands where there is some watertable ﬂuctuation and the surface becomes aerated by mid-season. These sites are often hummocked, with shrubs rooting on elevated microsites. Betula nana and Carex aquatilis are the characteristic species but Salix pedicellaris and Carex utriculata dominate on wetter sites. The moss layer is variable and can be diverse, absent, or dominated by Tomentypnum nitens, Sphag- num, or Drepanocladus. Some drier sites will have scattered, stunted trees (spruce or black spruce most commonly). Common soil types are terric and typic Mesisols and Fibrisols. Peat depths are frequently between 1 and 2 m but deep sedge-derived peat to 4 m occurs; this Site Association can occassionally occur on thin organic veneers.

Characteristic Vegetation Wetland Edatopic Grid

Tree layer (0 - 0 - 10) Soil Nutrient Regime Shrub layer (10 - 35 - 100) ABCD E F Betula nana,Salix pedicellaris Herb layer (5 - 60 - 100) VM Carex aquatilis,C. utriculata, Comarum palustre W Moss layer (0 - 70 - 100) AkVA N MA SA Aulacomnium palustre,Drepanocladus aduncus,Sphagnum Group I, VW

Tomentypnum nitens Soil Moisture Regime pH

Comments St Sl Mo Dy VD Hydrodynamic Index The Wf02 Site Association often occurs around the periphery of the wetter Wf01 or adjacent to the drier Wb05 .These three Site Associations may represent a sequence of long-term peatland succession.Many sites have a moss layer with rich and poor site indicators,suggesting that they are in transition from fen to bog conditions. The Wf02 is one of the most common Interior peatland community types at low to subalpine elevations.It is probably only absent from the AT,BG,and PP zones.In coastal areas,similar sites are occupied by the Wf52.

Chapter 5.2 Fens 83 Wf03 Water sedge – Peat-moss

Carex aquatilis – Sphagnum

General Description Water sedge – Peat-moss fens occur mainly at elevations above 1100 m in the Interior (ESSF zone), where they are the counterpart to the Wf02 of lower elevations. These communities appear to be relatively common but have not been extensively sampled. Small pocket depressions or gradual seepage slopes where there is no ﬂood- ing are typical locations. Carex aquatilis is the dominant species, though there can be signiﬁcant occurrence of subalpine forbs such as Caltha leptosepala, Sanguisorba canadensis, or Senecio triangularis on some sites. Peat-mosses are usually dominant in the Wf03, though there may be a diversity of other mosses such as Aulacomnium palustre, Tomentypnum nitens, and others. Mesisols derived from sedge peat up to 2 m (rarely to 4 m) in depth are common.

Characteristic Vegetation Wetland Edatopic Grid Tree layer (0 - 0 - 0) Shrub layer (0 - 3 - 10) Soil Nutrient Regime Herb layer (25 - 70 - 100) ABCD E F Carex aquatilis,C.sitchensis, VM Senecio triangularis Moss layer (30 - 85 - 100) W Aulacomnium palustre,Sphagnum Group I, AM ANAkVA N MA SA Tomentypnum nitens

VW Comments Soil Moisture Regime pH The Wf11 and Wf12 occur only at higher

elevations but require greater surface water- St Sl Mo Dy VD ﬂow and replace the Wf03 on active seeps Hydrodynamic Index and more saturated sites.Frost and cold soils rather than a high watertable probably limit shrub establishment on Wf03 sites.

84 Wetlands of British Columbia: A Field Guide to Identiﬁcation Barclay’s willow – Water sedge – Glow moss Wf04

Salix barclayi – Carex aquatilis – Aulacomnium palustre

General Description Barclay’s willow – Water sedge – Glow moss fen/swamps are common at subalpine elevations of the Sub-Boreal Interior, Southern Interior Mountains, and Northern Boreal Moun- tains. They occur on subalpine seepage slopes, along glacier-fed creeks, and in frost-prone basins. Salix barclayi dominates the shrub layer with a scattering of other low shrub species. Carex aquatilis dominates the herb layer but is often accompanied by scattered high-elevation species such as Caltha leptosepala, Eriophorum angustifolium, and Leptarrhena pyrolifolia. The moss layer can be absent or moderately well developed. Continuous (often copious) groundwater or snowmelt seepage is typical, and soils are cold. Peat is often shallow because of low biomass production but occasionally deep sedge peat deposits are encountered. Common soil types include terric Mesisols, Humisols, and Fibrisols

Characteristic Vegetation Wetland Edatopic Grid Tree layer (0 - .5 - 3) Shrub layer (10 - 35 - 95) Soil Nutrient Regime Salix barclayi ABCD E F Herb layer (26 - 65 - 99) VM Calamagrostis canadensis,Carex aquatilis, C.sitchensis W Moss layer (0 - 15 - 95) Aulacomnium palustre,Mnium spp., AkVA N MA SA Philonotis fontana VW

Comments Soil Moisture Regime pH

Wf04 can occur alone or surrounding sedge St Sl Mo Dy VD or cotton-grass fens (Wf03 or Wf12),or in Hydrodynamic Index wet depressions within forb-rich subalpine meadows or carrs.The similar Sc03 is also common at high elevations in the Interior. However,the Sc03’s low shrub physiognomy is the result of cold-air drainage not wet soils, and it is characterized by subalpine forbs with few hydrophytes.

Chapter 5.2 Fens 85 Wf05 Slender sedge – Common hook-moss

Carex lasiocarpa – Drepanocladus aduncus

General Description Slender sedge – Common hook-moss fens are common throughout the Interior at elevations below 1400 m. These fens occur on peat flats surrounding small lakes and ponds or in infilled palustrine basins. Pro- longed shallow surface flooding and continual surface peat saturation are typical. Carex lasiocarpa and Drepanocladus aduncus are constant dominants. Other large water sedges, such as C. aquatilis and C. utriculata, are also common. There can be a very sparse shrub cover of Salix pedicellaris, S. candida, or Betula nana. The moss layer is usually well developed but is occasionally absent. Hook- mosses usually dominate with occasional inclusions of other brown mosses. Deep peat deposits are common but some sites may occur on thin organic veneers. Mesisols are the most common soil type but Humisols and Fibrisols also occur.

Characteristic Vegetation Wetland Edatopic Grid Tree layer (0 - 0 - 0) Shrub layer (0 - 3 - 10) Soil Nutrient Regime Herb layer (13 - 60 - 100) ABCD E F Carex aquatilis,C. lasiocarpa,C.utriculata VM Moss layer (0 - 55 - 100) Drepanocladus aduncus W AM ANAkVA N MA SA Comments Some Wf05 sites are marsh-like with deep VW

ﬂooding,low diversity,and virtually no moss Soil Moisture Regime pH layer.The related Wf06 occurs on ﬂoating

mats with a more equable water regime and St Sl Mo Dy VD hummock/hollow topography.Slender- Hydrodynamic Index sedge fens (Wf05,Wf06) occur in locations similar to the Wf01 but seem to represent sites with longer surface saturation and more basic soil water.Similar sites in coastal areas are described by the Wf53.

86 Wetlands of British Columbia: A Field Guide to Identiﬁcation Slender sedge – Buckbean Wf06

Carex lasiocarpa – Menyanthes trifoliata

General Description Slender sedge – Buckbean fens are uncommon in the Central and Sub-Boreal Interior at elevations below 1300 m. They occur on ﬂoating mats adjacent to small lakes and peatland ponds, or in ﬂarks of patterned fens where there is permanent surface saturation and shallow inundation. Sites are often slightly hummocked, with Menyanthes trifoliata occurring in the wet depressions and Carex lasiocarpa and Drepanocladus spp. and other mosses occurring on mounds. A sparse shrub layer can occur and the moss layer is always well developed. Hook-mosses are the most common component of the moss layer but Sphagnum spp. or Campylium stellatum may dominate on some sites. Sites have sedge peat to 2.5 m, often with a subsurface water lens or su- persaturated horizon. Fibrisols and Mesisols are typical soil types.

Characteristic Vegetation Wetland Edatopic Grid Tree layer (0 - 0 - 0) Shrub layer (0 - 3 - 10) Soil Nutrient Regime Herb layer (20 - 65 - 85) ABCD E F Carex lasiocarpa, Menyanthes trifoliata VM Moss layer (30 - 55 - 100) Drepanocladus aduncus,Warnstorﬁa spp. W AM ANAkVA N MA SA Comments Wf06 site conditions are intermediate be- VW tween the Wf05 and the Wf08.The Wf06 Soil Moisture Regime pH has a more equable water regime with less

flooding,less water flow,and greater peat St Sl Mo Dy VD saturation than the related Wf05,but has Hydrodynamic Index deeper and more dynamic surface water than the Wf08.Slender-sedge fens (Wf05,Wf06) occur in locations similar to the Wf01 but seem to represent sites with longer surface saturation and more basic soil water.The Wf06 almost always occurs as a floating mat adjacent to a waterbody. Similar sites in coastal areas are described by the Wf53.

Chapter 5.2 Fens 87 Wf07 Scrub birch – Buckbean – Shore sedge

Betula nana – Menyanthes trifoliata – Carex limosa

General Description Scrub birch – Buckbean – Shore sedge fens occur throughout the Central and Sub-Boreal Interior at middle elevations below 1400 m, in palustrine basins or patterned fens with permanently high watertables. Most sites are prominently hummocked or ribbed with elevated sites and permanent shallow-water hollows. An open cover of Betula nana or Salix pedicellaris rooted on elevated microsites is distinctive. Low sedges such as Carex chordorrhiza and C. limosa are prominent throughout most sites, while Comarum palustre and Menyanthes trifoliata occupy inundated depressions. The composition of the well-developed bryophyte layer is variable. Mixed-species Sphagnum cover is common on some sites (not necessarily the most acidic), brown mosses are common on others, while true calciphiles such as Scorpidi- um scorpioides occur only on the most basic sites. Mesisols and Fibrisols derived from sedge/moss peat are typical. Peat is often >1 m in depth but, less commonly, sites occur on peat veneers.

Characteristic Vegetation Wetland Edatopic Grid Tree layer (0 - 0 - 0) Shrub layer (10 - 26 - 55) Soil Nutrient Regime Betula nana,Salix pedicellaris ABCD E F Herb layer (20 - 60 - 80) VM Carex aquatilis,C.chordorrhiza,C. limosa, Comarum palustre,Menyanthes trifoliata W Moss layer (20 - 95 - 100) Drepanocladus aduncus,Scorpidium spp., AkVA N MA SA Sphagnum Group I,Sphagnum Group II, Tomentypnum nitens VW

Soil Moisture Regime pH Comments St Sl Mo Dy VD The Wf07 covers much of the acidity/alkalini- Hydrodynamic Index ty gradient,with little change in the vascular ﬂora but a marked difference in the bryophyte composition.Some sites have more Sphagnum and are generally more bog-like, while others are more clearly rich fens.These sites are similar to the Wb13 but have a well developed shrub layer.However,there is a high degree of intergradation with few clear environmental criteria to separate most sites (except at the extremes of the spectrum).Hence,the variation has been grouped into a single Site Association. The open shrub cover of the the Wf07 distinguishes it from the wetter Wf08 or Wb13.The Wf07 may represent the middle stage of a peatland succession sequence: Wf08 >> Wf07 >> Wb11 in some regions .

88 Wetlands of British Columbia: A Field Guide to Identiﬁcation Shore sedge – Buckbean – Hook-moss Wf08

Carex limosa – Menyanthes trifoliata – Drepanocladus

General Description The Shore sedge – Buckbean – Hook-moss is an uncommon, rich Fen Site Association that occurs mainly at higher elevations throughout the Interior (700–1800 m) in colder subzones. These fens occur on pond-side floating mats or in flarks of patterned fens where there is prolonged shallow flooding to no more than several centimetres. Carex limosa rooted in shallow water is the constant dominant on these sites. Menyanthes trifoliata occurs on most sites but can be very sparse or absent on some. A diversity of species tolerant of permanent saturation such as Carex chordorrhiza, Equisetum fluviatile, and Andromeda polifolia commonly occur with low cover. Peat deposits are shallow (0.5 m) to very deep (> 6 m), fibric or mesic, and derived from fine sedges and brown mosses. Fibrisols are the most common soil type.

Characteristic Vegetation Wetland Edatopic Grid Tree layer (0 - 0 - 0) Shrub layer (0 - .5 - 10) Soil Nutrient Regime Herb layer (14 - 35 - 100) ABCD E F C. limosa,Menyanthes trifoliata VM Moss layer (1 - 85 - 100) Drepanocladus spp. W AM ANAkVA N MA SA Comments This is the most common and dominant Site VW

Association in patterned fens.In weakly pat- Soil Moisture Regime pH terned fens,the Wf08 occurs over ribs and

flarks.Where there is a more pronounced St Sl Mo Dy VD rib/flark pattern,the Wf08 will typically Hydrodynamic Index occur in flarks and the floristically similar, shrubby Wf07 on elevated ribs. The Wf06 occurs on wetter and more hydrologically dynamic sites than the Wf08.Similarly stagnant sites with acidic soil water are occupied by the Wb13.The Wf08 has similar hydrology to the Wf09 and Wf10,but with more mobile groundwater and greater degree of surface flooding.Wf08 sites may become Wb13 sites in some circumstances. Peat deposits are often consistent throughout the profile,and peat core contents of fine sedge and brown mosses are readily identifiable.This suggests that these ecosystems can be stable and long-lived.

Chapter 5.2 Fens 89 Wf09 Few-ﬂowered spike-rush – Hook-moss

Eleocharis quinqueﬂora – Drepanocladus

General Description The Few-flowered spike-rush – Hook-moss Fen Site Associa- tion occurs on small sloping peatlands at high elevations (mostly above 1200 m) throughout the Sub-Boreal, Central, and South- ern Interior. It is rare throughout most of its range, occurring only in slope positions with continual slow surface seepage. Plant diversity is low; Eleocharis quinqueflora is the site dominant, with lesser amounts of Carex limosa, Eriophorum angustifolium, and other forbs occasionally occurring. Hook- mosses such as Homatocaulis vernicosus, Scorpidium revolvens, and Drepanocladus aduncus usually comprise the moss layer but other brown mosses such as Meesia triquetra and Tomentypnum nitens can occur in high abundance. Peat forms as a characteristically dense and tenacious mesic peat. Peat depths are frequently shallow but can be up to 2 m. Terric Mesisols and Humisols are common soil types.

Characteristic Vegetation Wetland Edatopic Grid Tree layer (0 - 0 - 0) Shrub layer (0 - .5 - 10) Soil Nutrient Regime Herb layer (30 - 60 - 100) ABCD E F Carex limosa,Eleocharis quinqueﬂora, VM Eriophorum angustifolium Moss layer (1 - 50 - 95) W Drepanocladus spp.,Tomentypnum nitens AM ANAkVA N MA SA

Comments VW

The Wf09 unit is similar in structure and hy- Soil Moisture Regime pH droedatopic position to the Wf11 of lower

elevations.Wf09 commonly occurs without St Sl Mo Dy VD adjacent wetland Site Associations or in Hydrodynamic Index complex with cotton-grass fens (Wf12 or Wf13). The tenacious peat of this unit is typically of similar composition throughout the profile, suggesting that this ecosystem can be stable and long-lived. Peat is sufficiently dense on Wf09 sites that soil water movements are impeded and most waterflow is at the surface as sheet flow.The specific conditions that give rise to the Wf09 rather than other high-elevation fens are not well understood but may be partly initiated and maintained by the dense stems and roots of Eleocharis quinqueflora.

90 Wetlands of British Columbia: A Field Guide to Identiﬁcation Hudson Bay clubrush – Red hook-moss Wf10

Trichophorum alpinum – Scorpidium revolvens

General Description The Hudson Bay clubrush – Red hook-moss Site Association is rare and seems to occur only in the moist subzones of the SBS. It is floristically related to the Wf11 but occurs where the watertable is more stagnant and at or slightly above the peat surface for much of the growing season. Com- mon locations are around small peatland lakes and ponds, and in flarks. Trichophorum alpinum dominates but there is usually a diverse array of other species typical of base-rich and saturated peatland habitats such as Carex chordorrhiza, C. lasiocarpa, C. limosa, Menyanthes trifoliata, and Triglochin maritima. The moss flora is dominated by brown mosses and strong calciphiles: Campylium stellatum, Scorpidium revolvens, and S. scorpioides. Many sites are underlain by calcareous marl, and peat water is usually neutral to alkaline. Peat decomposition is apparently retarded under these conditions because the entire peat proﬁle often has few signs of decomposition. Typic Fibrisols are common. Peat is usually deep (> 3 m).

Characteristic Vegetation Wetland Edatopic Grid Tree layer (0 - 0 - 0) Shrub layer (0 - 0 - 0.5) Soil Nutrient Regime Herb layer (37 - 90 - 95) ABCD E F Andromeda polifolia,Carex aquatilis, VM C. chordorrhiza,C.lasiocarpa,C. limosa, Drosera anglica,Menyanthes trifoliata, Oxycoccus oxycoccos,Triantha glutinosa, W Trichophorum alpinum,Triglochin mariti- AkVA N MA SA ma Moss layer (25 - 95 - 100) VW

Calliergon stramineum,Campylium stella- Soil Moisture Regime pH tum,Scorpidium revolvens,S. scorpioides,

Sphagnum Group II,Tomentypnum nitens St Sl Mo Dy VD Hydrodynamic Index Comments Wf10 sites are usually small inclusions in larger complexes of rich fen peatland but are conspic- uous when Trichophorum alpinum is in ﬂower.Most site conditions are similar to the Wf08 but the Wf10 occurs where soil water is alkaline and (apparently) occurs over a much more restricted climatic range.

Chapter 5.2 Fens 91 Wf11 Tufted clubrush – Star moss

Trichophorum cespitosum – Campylium stellatum

General Description The Tufted clubrush – Star moss Fen Site Association is scattered throughout the Interior at middle to subalpine elevations, most commonly in regions underlain with base-rich parent materials. These fens occur on level and gently sloping, groundwater-fed peatlands that are permanently saturated but rarely inundated. Sites have smooth, ribbed, or slightly hummocked topography and any depressions are water-ﬁlled. Trichophorum cespitosum and Campylium stellatum are constant dominants and occur mainly on drier microsites. Menyanthes trifoliata and calcium-encrusted Scorpidium scorpioides and Scorpidium revolvens are commonly found in very shallow pools. Most sites have a distinct dense and tenacious turfy peat. Deep peat is typical (to 5 m) but occasionally thin peat veneers occur. Fibrisols and Mesisols are typical soil types.

Characteristic Vegetation Wetland Edatopic Grid

Tree layer (0 - 0 - 0) Soil Nutrient Regime Shrub layer (0 - 1 - 10) ABCD E F Herb layer (20 - 75 - 97) Carex limosa,Eriophorum angustifolium, VM Menyanthes trifoliata,Trichophorum cespitosum W Moss layer (0 - 70 - 95) AkVA N MA SA Campylium stellatum,Sphagnum Group II VW

Comments Soil Moisture Regime pH The Wf11 occurs where extremely high pH St Sl Mo Dy VD limits the availability of phosphorous,mak- Hydrodynamic Index ing these sites nutrient-poor even though they have an abundance of cations.Tufted clubrush–dominated wetlands are also found in regions underlain by base-poor granitic parent material,such as coastal British Columbia,where phosphorus is also limited.These communities lack minerotrophic site indicators and have a Sphagnum-dominated moss layer. Tufted clubrush – Peat-moss ecosystems (Wb52) are very common in coastal British Columbia but several sites have been observed in interior locations where the local geology is of igneous intrusive origin (e.g.,Monashee Ranges).

92 Wetlands of British Columbia: A Field Guide to Identiﬁcation Narrow-leaved cotton-grass – Marsh-marigold Wf12

Eriophorum angustifolium – Caltha leptosepala

General Description The Narrow-leaved cotton-grass – Marsh-marigold Site Asso- ciation is common at subalpine elevations (above 1200 m) throughout the Sub-Boreal and Central Interior. It occurs on gently sloping peatlands where there is continual seepage from snowmelt and groundwater. Eriophorum angustifolium occurs on most sites with high cover. Sites with abundant surface seepage will also have a high cover of Caltha leptosepala and/or Leptarrhena pyrolifolia. Other graminoids such as C. anthoxanthea, C.aquatilis, or C.nigricans may also occur with high cover on some sites. The moss layer is usually well developed but compositionally variable. Soils are usually deep, mushy sedge peat. Typic Mesisols and Fibrisols are the most common soil types.

Characteristic Vegetation Wetland Edatopic Grid Tree layer (0 - 0 - 0) Shrub layer (0 - 1 - 10) Soil Nutrient Regime Herb layer (12 - 80 - 100) ABCD E F Caltha leptosepala,Eriophorum angusti- VM folium Moss layer (0 - 75 - 95) W Aulacomnium palustre AM ANAkVA N MA SA

Comments VW

The Wf12 occurs on sites with more active Soil Moisture Regime pH seepage than the related Wf13 Site Associa- tion.It also has similar site characteristics to St Sl Mo Dy VD the Wf08,but that unit is fed by groundwater Hydrodynamic Index with high levels of base cations and has dense peat deposits. The Wf12 occurs alone or in complex with the Wf03,on microsites with more active seepage. Some Wf12 sites in the upper Skeena drainage have high cover of Carex anthoxanthea,which is a common species of bog forests on the north Coast.The Interior distribution of this species is greatly restricted and could be limited to these high-elevation wetland ecosystems.

Chapter 5.2 Fens 93 Wf13 Narrow-leaved cotton-grass – Shore sedge

Eriophorum angustifolium – Carex limosa

General Description Narrow-leaved cotton-grass – Shore sedge fens occur at higher elevations (1200–1800 m) of the ESSF zone in depressions or gradual seepage slopes where standing water persists for most of the short growing season. The Wf13 appears to be relatively common (at least locally) but has not been extensively sampled. A community dominated by Eriophorum angustifolium with Carex limosa is typical but some sites may have poor sedge (Carex magellanica) instead of C. limosa. Grasses such as Calamagrostis canadensis and Vahlodea atropurpurea and the forb Caltha leptosepala are commonly abundant. The moss layer is well developed and is often diverse, with no one species dominating. Soils are deep peat deposits of ﬁbric or mesic cotton-grass remains. Typic Mesisols and Fibrisols are common soil types.

Characteristic Vegetation Wetland Edatopic Grid Tree layer (0 - 0 - 0) Shrub layer (0 - 2 - 10) Soil Nutrient Regime Herb layer (20 - 80 - 100) ABCD E F Caltha leptosepala,Carex aquatilis, VM C. limosa,Eriophorum angustifolium Moss layer (0 - 40 - 99) W Aulacomnium palustre,Philonotis fontana, Sphagnum Group I AkVA N MA SA

VW Comments Soil Moisture Regime pH The Wf13 is wetter than the closely related

Wf12 and tends to be found more commonly St Sl Mo Dy VD in depressional areas where water ponds. Hydrodynamic Index The high-elevation fen units Wf03,Wf04, Wf11,Wf12,and Wf13 often occur together in complex in extensive subalpine peatlands, each occurring in habitats differing in water ﬂow and ponding (Wf03 driest to Wf13 wettest).

94 Wetlands of British Columbia: A Field Guide to Identiﬁcation Narrow-leaved cotton-grass – Peat-moss Wf50

Eriophorum angustifolium – Sphagnum

General Description The Narrow-leaved cotton-grass – Peat-moss Fen/Bog Site Association describes a wide range of ecosystems of montane and subalpine areas of the Coast where there is some surface seepage. Many sites are sloping but the Wf 50 also occurs on level sites with a permanent, high watertable. As with many sloping peatlands, sites are microtopographically heterogeneous and therefore a mosaic of vegetation is common. Eriophorum angustifolium is always prominent but other species can be very abundant on some sites or in speciﬁc locations within the peatland. Tree and shrub species, if they occur, are on raised sites. Groundwater seepage pools or surface drainage channels are common. Peat deposits are generally < 2 m deep. Mesic sedge peat throughout the proﬁle is common. Terric and Typic Mesisols are common soil types.

Characteristic Vegetation Wetland Edatopic Grid Tree layer (0 - 0 - 0) Shrub layer (0 - 3 - 10) Soil Nutrient Regime Herb layer (39 - 75 - 98) ABCD E F Eriophorum angustifolium,Fauria VM crista-galli,Kalmia microphylla, Trichophorum cespitosum W Moss layer (20 - 75 - 99) AM ANAkVA N MA SA Sphagnum Group I

VW Comments Soil Moisture Regime pH The Wf50 is analogous to the interior Wf11 and Wf12 of subalpine sloping seeps but has St Sl Mo Dy VD many coastal species and is generally more Hydrodynamic Index heterogeneous.The complex nature of hydrological flow in these sites means that large tracts of homogeneous vegetation are uncommon.Several more specific classification units based on the presence of additional dominants are likely possible with sufficient data.Sites with high cover of Carex pauciflora,Sanguisorba spp.,Fauria crista-galli,Dodecatheon jeffreyi,or Rubus chamaemorus have all been observed.

Chapter 5.2 Fens 95 Wf51 Sitka sedge – Peat-moss

Carex sitchensis – Sphagnum

General Description Sitka sedge – Peat-moss fens occur at low elevations along the Coast, in wet drainage channels or hollows in sloping peatlands where there is gradually ﬂowing surface water. These sites are uncommon and often of small areal extent relative to other ecosystems of the Coast. Carex sitchensis grows in dense swards with Sphagnum species in carpets or ﬂoating in shallow water. A diversity of other species occurs with low cover on most sites. Peat accumulations in the Wf 51 range from thin veneers to deep blankets of poorly to well-decomposed peat. Organic layers are often intermixed with mineral materials. Fibrisols are the most common soil type but Mesisols and Hu- misols also occur.

Characteristic Vegetation Wetland Edatopic Grid Tree layer (0 - 0 - 0) Shrub layer (0 - .5 - 5) Soil Nutrient Regime Herb layer (15 - 82 - 100) ABCD E F Carex sitchensis,Comarum palustre VM Moss layer (0 - 40 - 100) Sphagnum spp. W AM ANAkVA N MA SA Comments The Wf51 is similar to the Wf01 in most re- VW

spects but has a coastal distribution; Carex Soil Moisture Regime pH sitchensis replaces Carex aquatilis on the

Coast.The presence of Sphagnum on these St Sl Mo Dy VD sites reﬂects the wider tolerance to variable Hydrodynamic Index hydrology of Sphagnum species that occur in this climate and not to ombrotrophic conditions,which is typical in the Interior. The Wm50 occurs on more hydrologically active sites than the Wf51.

96 Wetlands of British Columbia: A Field Guide to Identiﬁcation Sweet gale – Sitka sedge Wf52

Myrica gale – Carex sitchensis

General Description Sweet gale – Sitka sedge fens are uncommon at low elevations in the Georgia Depression and Coast and Mountains in a wide variety of landscape positions. Sites can be shallowly ﬂooded in the early season but will drop just below the surface for most of the growing season. Myrica gale and Spiraea douglasii form a closed and sometimes dense thicket mostly < 1.5 m in height. Carex sitchensis dominates the herb layer but there is a scattering of other species on most sites. Be- cause of ﬂooding, the bryophyte layer is generally sparse but on some sites cover of Sphagnum or other moss species may be high. Peat deposits are mostly shallow, moderately to well decomposed sedge and wood peat. Terric Humisols and Mesisols are common soil types.

Characteristic Vegetation Wetland Edatopic Grid Tree layer (0 - 0 - 0) Shrub layer (30 - 60 - 85) Soil Nutrient Regime Myrica gale,Spiraea douglasii ABCD E F Herb layer (20 - 35 - 100) VM Carex sitchensis Moss layer (0 - 14 - 60) W Sphagnum spp. AM ANAkVA N MA SA

Comments VW

The Wf52 Site Association is common as a Soil Moisture Regime pH component of many peatlands in the south

Coast.It is most often found in complex with St Sl Mo Dy VD the Wm50 in more peripheral (and drier) Hydrodynamic Index locations but occurs around other Site Associ- ations as well,including estuarine marshes. Shrub thickets dominated by Spiraea douglasii with sparse Myrica gale and Carex spp. are common in the region where the Wf52 occurs.These communities are usually on mineral soil and described by the Ws50 Site Association.

Chapter 5.2 Fens 97 Wf53 Slender sedge – White beak-rush

Carex lasiocarpa – Rhynchospora alba

General Description Slender sedge – White beak-rush fens occur in the Georgia Depression at elevations below 600 m. The Wf 53 requires permanently saturated soils and is tolerant of prolonged shallow inundation. Lake margins are the most common location but some isolated basins may also have suitable conditions. Carex lasiocarpa is always abundant and dominant. Rhynchospora alba and scattered low-growing Myrica gale occur on most sites. However, wetter sites often lack these species and have aquatic species such as Nuphar lutea, Brasenia schreberi, or Menyanthes trifoliata instead. Schoenoplectus acutus grows in more alkaline examples of the Wf 53. Soils are shallow mesic or humic peat of sedge or limnic origin. Terric Mesisols and Humisols are common soil types.

Characteristic Vegetation Wetland Edatopic Grid Tree layer (0 - 0 - 0) Shrub layer (0 - 3 - 10) Soil Nutrient Regime Myrica gale ABCD E F Herb layer (60 - 80 - 85) VM Carex lasiocarpa,Rhynchospora alba Moss layer (0 - 2 - 8) W AM ANAkVA N MA SA Comments Coastal Carex lasiocarpa stands occur on a VW

range of ecological conditions from semi-ter- Soil Moisture Regime pH restrial to shallowly ﬂooded and marsh-like

peatlands.Ceska (1978) observed variants of St Sl Mo Dy VD this unit: a typic with Rhynchospora alba,a Hydrodynamic Index limose variant with Nuphar lutea,a mineric variant with Dulichium arundinaceum and Schoenoplectus acutus,and a higher- elevation variant with Carex limosa.With sufficient additional data,several more specific classification units based on the presence of additional dominants might be indicated.Carex lasiocarpa communities with abundant Spiraea douglasii and Myrica gale occur on drier and hummocky sites (see additional units). The Wf52 is similar but grows on drier and more acidic sites than the Wf53.The Wf53 is analogous to the Wf05 of the Interior but has little or no moss cover and includes coastal species.

98 Wetlands of British Columbia: A Field Guide to Identiﬁcation Additional Fen Site Associations

Sweet gale – Pink spirea – Slender sedge Myrica gale – Spiraea douglasii – Carex lasiocarpa Sweet gale – Pink spirea – Slender sedge fens are uncommon in the Georgia Depression in basins and lake margins. The shrub layer is well developed and composed of Myrica gale and/or Spireae douglasii. Carex lasiocarpa is dominant in the herb layer but a diversity of other herbs occurs, including Carex sitchensis, Hypericum anagalloides, Menyanthes trifoliata, and Comarum palustre. The moss layer is very sparse or absent. Soils are primarily Mesisols. These communities occur on sites drier or with more microtopography than the Wf 53. Similar but more acidic sites support the Wf 52.

Menyanthes trifoliata, buckbean

Chapter 5.2 Fens 99 1

1 Carex chordorrhiza, cordroot sedge 2 Carex lasiocarpa, slender sedge 3 Eriophorum angustifolium, narrow-leaved cotton-grass

100 Wetlands of British Columbia: A Field Guide to Identiﬁcation 5.3 MARSHES

1 A pothole marsh dominated by the rare grass Scolochloa festucacea, the Wineglass Ranch near Riske Creek (IDFdk4) 2 A coastal Beaked sedge marsh on Minerva Lake near Prince Rupert (CWHvh2) 3 Cattail thicket, Nicola Lake (BGxw1)

Chapter 5.3 Marshes 101 MARSH WETLAND CLASS

Definition A marsh is a permanently to seasonally flooded non-tidal mineral wetland dominated by emergent grass-like vegetation. General Description Vegetation Table 5.3.2 lists species common in the Marsh Site Associations described in this guide. Marshes are floristically simple plant communities with low species diversity and strong dominance by one or two species. The high nutrient availability in marshes favours “aggressive” species that spread vegetatively. This results in communities with one or two dominant species that effectively limit establishment and spread of other species. Dominance can result from optimal environmental conditions for mature plants, favourable conditions for initial establishment of one species over another, or simply chance initial establishment of one species. Marshes have > 10% cover of emergent grasses, rushes, sedges, or (occasionally) forbs or horsetails. The tree, shrub, and bryophyte layers in marshes are usually absent or very sparse (< 10%). Aquatic plants are common, especially in marshes that retain standing water for most or all of the year. Landscape Position and Distribution Marshes are favoured by dynamic hydrological regimes, high nutrient status, and warm growing-season climates. In cool, wet climates, marshes are restricted to wave-washed lakeshores, stream floodplains, and back-levees where waterflow prevents peat accumulation and keeps nutrient availability high. In these climates, peatlands occupy most small basins. In warm and dry climates, however, marshes are the most common wetland class and occur in most hydrogeomorphic types including small potholes and depressions. In these areas, high evapotranspiration rates result in watertable fluctuations that expose the soil surface in late season and promote decomposition of organic materials. Large variation in pothole water levels between years is common in semi-arid climates. The Site Associations and their extent often varies considerably in response to these hydrological changes.

102 Wetlands of British Columbia: A Field Guide to Identification Hydrology and Soils Marshes are always flooded in the Soil Nutrient Regime early season to depths up to 3 m. ABCD E F Some remain flooded throughout M the year and have circulating wa- VM ters that maintain a high nutrient availability (Figure 5.3.1). Howev- W AM ANAkVA N MA SA er, many have pronounced Marshes drawdown and substrate expo- VW sure by mid to late summer. Soils Soil Moisture Regime pH are usually mineral, but they can also have a well-decomposed or- St Sl Mo Dy VD Hydrodynamic Index ganic surface tier of humic or limnic peat. Nutrient availability  .. Position of marshes on the is high due to circum-neutral pH, edatopic grid. abundant waterflow, and periodic exposure and aeration of the substrate. Conservation Issues Marshes are critically important wetland ecosystems. They are the most heavily used wetland type for most wetland-using wildlife species because they support a large standing crop of palatable vegetation, plankton, and aquatic invertebrates—organisms that form a food base for larger animals. Marshes are the favoured wetland class for most waterfowl, amphibians, and semi-aquatic mammals because they provide good cover, open water, and a food source for young animals. They are the most common wetland type in the dry and warm climates where wetlands in general are uncommon and this is where many of British Columbia’s rare wetland-dependent vertebrates occur (e.g., Great Basin Spadefoot, Tiger Salamander). Marshes are early-seral ecosystems in wetland succession. Marshes are the easiest wetland class to create artificially because they will form naturally in recently created wetland environments (e.g., roadside ditches, sewage lagoons). Most marshes are tolerant of hydrological modifications that are not outside the natural, broad range for the Site Association. Maintaining wetland habitats by stabilizing watertables is a common practice in the prairie pothole region of Canada, where many wetlands dry up during drought years. However, throughout most of British Co- lumbia, this method is inappropriate and will reduce the productivity of existing marshes. Most marshes will recover from even severe mechani- cal or grazing disturbance if hydrological regime is maintained.

Chapter 5.3 Marshes 103 Phragmites australis (common reed), Lythrum salicaria (purple loosestrife), and Iris pseudacorus (yellow-ﬂag) are invasive wetland species in British Columbia. Like other marsh dominants, these require nutrient- rich site conditions and warm growing-season temperatures. However, these species require more growing degree days than occur in most of the province, thereby restricting them to warmer climates. It is unclear how profound the impact of invasive species is upon the rest of the ecosystem because they merely replace one near monoculture with another (Farnsworth and Ellis 2001; Gardner et al. 2001). Waste-water treatment wetlands will be marshes. High inputs of nitroge- nous waste will drive any wetland towards marsh-like conditions.

Lemna minor, duckweed

104 Wetlands of British Columbia: A Field Guide to Identiﬁcation s xx xx xxxxx BGPP BWBS SWB ESSF ICH IDF MS SBS CDF CWH SBPS MH Distribution of Marsh Site Associations by biogeoclimatic zone Beaked sedge – Water sedgeWater sedge – Beaked sedge horsetail – Beaked Swamp sedgeAwned spike-rushCommon Cattail bulrushGreat Baltic rush sedge – Hemlock-parsley Sitka sedgeThree-way x x xx x x xxx x xxx x xx xxx xx x xxx x xx x x x x x x xx xx xx xx x xx x x xx x xx x xx x x x x x = incidental; < 5% of wetlandss = southern subzones only xx = minor; 5–25% of wetlands xxx = major; >25% of wetlands Wm01 Wm02 Wm03 Wm04 Wm05 Wm06 Wm07 Wm50 Wm51  .. 

Chapter 5.3 Marshes 105  .. Marsh Species Importance Table

Species Wm01 Wm02 Wm03 Wm04 Wm05 Herbs Carex utriculata xyzzz xyzzz x xy and Carex aquatilis xyzz xy x Dwarf Equisetum fluviatile x xyzzzz x Shrubs Comarum palustre xy xy x Sium suave xxy Carex exsiccata x Carex atherodes xyzzzz Polygonum amphibium xxxyxx Eleocharis palustris x xyzzzz x Potamogeton richardsonii xyz Typha latifolia x xyzzzz Schoenoplectus acutus x Menyanthes trifoliata Utricularia macrorhiza xxxxx Juncus balticus xx Hordeum jubatum x Potentilla anserina x Calamagrostis canadensis xy x xy x Cicuta douglasii xx Lysichiton americanus x Oenanthe sarmentosa x Galium trifidum x Spiraea douglasii x Carex sitchensis Nuphar lutea ssp. polysepala xx Dulichium arundinaceum Mosses Drepanocladus spp. xy xy xy x x Warnstorfia spp. xxxxx

106 Wetlands of British Columbia: A Field Guide to Identification Wm06 Wm07 Wm50 Wm51 Common Name xxxx beaked sedge water sedge x swamp horsetail xy xy marsh cinquefoil hemlock water-parsnip xxxy inflated sedge x awned sedge x water smartweed xx common spike-rush Richardson’s pondweed xx common cattail xyzzzz x great bulrush xxybuckbean xxgreater bladderwort x xyzzzz Baltic rush xyz foxtail barley x xyz common silverweed xx bluejoint x Douglas’ water-hemlock xy skunk cabbage xyzz Pacific water-parsley xyz small bedstraw xy x pink spirea xyzzzz xyz Sitka sedge x x xyz yellow pond-lily xyzzzz three-way sedge x hook-mosses: intermediate xx hook-mosses: poor

Chapter 5.3 Marshes 107 Wm01 Beaked sedge – Water sedge

Carex utriculata – Carex aquatilis

General Description Beaked sedge – Water sedge marshes constitute the most common and widespread Marsh Site Association in the province. The Wm01 occurs in all subzones from low to subalpine elevations on sites that are inundated by shallow, low-energy floodwaters and that experience some late-season drawdown. These marshes are found in a wide variety of landscape positions including flooded beaver ponds, lake margins, floodplains, and palustrine basins. Species diversity is low and plant cover is strongly dominated by Carex utriculata and C. aquatilis with scattered forbs, aquatics, and mosses. On sites experiencing significant surface drying, species diversity increases and sites become more meadow-like. Species such as Cala- magrostis canadensis, Geum macrophyllum, or Deschampsia cespitosa can become prominent. The Wm01 occurs over a wide range of site conditions on mineral substrates with thin peat veneers. Common soil types include Gleysols and Terric Humisols.

Characteristic Vegetation Wetland Edatopic Grid

Tree layer (0 - 0 - 0) Soil Nutrient Regime Shrub layer (0 - 0 - 5) ABCD E F Herb layer (13 - 80 - 100) Carex aquatilis, C.utriculata VM Moss layer (0 - 5 - 100) W Comments AkVA N MA SA The Wf01 and Wm01 have similar plant com- VW munities,but,because these units are species-poor and the two dominant sedge Soil Moisture Regime pH species have a wide ecological amplitude,the plant community poorly differentiates between St Sl Mo Dy VD Hydrodynamic Index sites on peat (Wf01) and those on mineral soil (Wm01).In general,the Wm01 is more deeply ﬂooded,has more dynamic hydrology,and has a higher cover of C. utriculata. The Wm02 is another similar community that occurs on more hydrologically dynamic locations such as lake margins or ﬂoodplains.In cooler climates the Wm01 frequently develops into Wf01 on sites with less dynamic hydrology. Some Wm01 sites have scattered tall shrubs; those sites supporting > 10% shrub cover are described by Swamp Site Associations (Section 5.4).

108 Wetlands of British Columbia: A Field Guide to Identiﬁcation Swamp horsetail – Beaked sedge Wm02

Equisetum ﬂuviatile – Carex utriculata

General Description The Swamp horsetail – Beaked sedge Marsh Site Association is uncommon at lower elevations throughout the Interior. Com- mon locations are in back-levee depressions along sediment- laden, low-gradient streams, protected bays of large lakes, or hydrologically modified (flooded) fens. The Wm02 also occurs along the Coast in tidal reaches of large rivers above saltwater influence. Plant diversity is low. Sites are dominated by Equisetum fluviatile with Carex utriculata sometimes co-dominating; often there are scattered aquatics such as Potamoge- ton and Myriophyllum spp. The Wm02 is similar to the Wm01 but is distinguished by its higher hydrodynamic index and by the dominance of E. fluviatile. Soils are derived from silty or fine-sandy fluvi- um, deep limnic deposits at open margins of lakes, or recently flooded peat. Rego Gleysols and Terric Humisols are common soil types.

Characteristic Vegetation Wetland Edatopic Grid

Tree layer (0 - 0 - 0) Soil Nutrient Regime Shrub layer (0 - 0 - 4) ABCD E F Herb layer (18 - 85 - 100) C. utriculata, Equisetum ﬂuviatile VM Moss layer (0 - 0 - 90) W Comments AkVA N MA SA

E. ﬂuviatile is tolerant of extreme variations VW in water depth and high rates of sedimentation and can colonize exposed mineral or Soil Moisture Regime pH peat soils.It has been used to revegetate the extreme environment of the drawdown zone St Sl Mo Dy VD Hydrodynamic Index in reservoirs. On ﬂuvial sites,the Wm02 is usually adjacent to tall-willow swamps or low bench communities.In lake systems,Wm02 commonly adjoins open water and other marsh communities.

Chapter 5.3 Marshes 109 Wm03 Awned sedge

Carex atherodes

General Description The Awned sedge Marsh Site Association is uncommon and restricted to dry climates of the Central Interior at low to middle elevations. These marshes are generally small and occur most commonly in small potholes surrounded by forest, where water levels are shallow and relatively constant. Sites are always dominated by Carex atherodes, but infrequently other species, such as Drepanocladus aduncus, Myriophyllum verticillatum, Alopecurus aequalis, or Carex utriculata, occur in abundance. Standing water is slightly alkaline; rooting substrates are ﬁne-textured mineral or shallow sedge-derived peat. Common soil types are Humisols and Humic Gleysols.

Characteristic Vegetation Wetland Edatopic Grid

Tree layer (0 - 0 - 0) Soil Nutrient Regime Shrub layer (0 - 0 - 4) ABCD E F Herb layer (55 - 90 - 100) Carex atherodes VM Moss layer (0 - 0 - 95) Drepanocladus aduncus W AM ANAkVA N MA SA Comments VW This unit is similar to the Wm01 but is much more limited in distribution and seems to be Soil Moisture Regime pH favoured by more alkaline waters.In the BG zone,Woolly sedge marshes occur on sites St Sl Mo Dy VD similar to the Wm03 (see additional units). Hydrodynamic Index Wm03 sites often occupy entire basins but they are also found in small patches within some larger Wm01 or Wm08 marshes. The distribution of Wm03 is primarily within rangelands and C. atherodes is palatable; many sites experience some level of grazing pressure.

110 Wetlands of British Columbia: A Field Guide to Identiﬁcation Common spike-rush Wm04

Eleocharis palustris

General Description Common spike-rush marshes are widely distributed throughout the Interior at elevations below 1300 m. They occur along lakeshores, and as a zone in larger potholes, oxbows, and slow-moving rivers, where there is some weak waterflow or wave action. Sites are shallowly flooded in the early season in all locations; the watertable often drops to the surface in palustrine locations but is permanent in lacustrine or fluvial systems. Wm04 sites also occur in freshwater and brackish tidal reaches of large coastal rivers and estuaries. Plant diversity is low; Eleocharis palustris is often the only emergent species with signiﬁcant cover. In interior sites, submerged and floating aquatics can be common; in estuarine sites Carex lyngbyei is often present. Soils are typically sandy or gravelly with or without a thin organic veneer.

Characteristic Vegetation Wetland Edatopic Grid

Tree layer (0 - 0 - 0) Soil Nutrient Regime Shrub layer (0 - 0 - 0) ABCD E F Herb layer (10 - 70 - 100) Eleocharis palustris, VM Potamogeton richardsonii Moss layer (0 - 0 - 10) W AM ANAkVA N MA SA Comments VW Eleocharis palustris occurs commonly in a wide variety of wetland habitats,including Soil Moisture Regime pH alkaline and weakly saline marshes,rich fens,and estuarine marshes.However,the St Sl Mo Dy VD Hydrodynamic Index Wm04 describes only those sites where E. palustris dominates.Wm04 sites are generally more shallowly ﬂooded than Wm06 sites and better aerated than Wm05 sites. The Wm04 commonly occurs adjacent to Wm05,Wm06,and Wm07 Site Associations or shallow-water ecosystems.

Chapter 5.3 Marshes 111 Wm05 Cattail

Typha latifolia

General Description Cattail marshes are common throughout the Coast and Interi- or at low elevations in subzones with warm summers. They occur most commonly in protected lake embayments and potholes or even roadside ditches, where the surface substrate remains saturated for most of the growing season. Typha latifolia dominates, often with few other rooted plants present, especially where nutrient levels are high and T. latifolia growth profuse. Occasionally there is signiﬁcant cover of Carex utriculata, Schoenoplectus acutus, or Lemna spp. These sites often have organic veneers of well-decomposed, odiferous muck. Soil types can be Humisols or Humic Gleysols. Water depths may be up to 1 m in the spring but recede in late summer, sometimes to the surface.

Characteristic Vegetation Wetland Edatopic Grid

Tree layer (0 - 0 - 0) Soil Nutrient Regime Shrub layer (0 - 0 - 10) ABCD E F Herb layer (40 - 80 - 100) Typha latifolia VM Moss layer (0 - 0 - 90) W Comments AkVA N MA SA

Typha latifolia effectively turns high nutrient VW levels (N and P) into biomass and often dominates wetlands experiencing nutrient Soil Moisture Regime pH loading.Addition of agricultural or human St Sl Mo Dy VD waste to most wetlands will lead to an in- Hydrodynamic Index crease and eventual dominance by T. latifolia if climatic conditions are favourable.Initial T. latifolia establishment requires substrate exposure for seedling establishment and germination,though once established it spreads extensively by rhizomes so that large stands may consist of only a few individual plants. Similar sites with more dynamic hydrology or lower N and P are usually occupied by Wm06. Patches of S. acutus in Wm05 marshes can be a result of intensive grazing by Muskrat. S. acutus stores nutrients in the root mass and can more rapidly recover from removal of its stem than can T. latifolia.

112 Wetlands of British Columbia: A Field Guide to Identiﬁcation Great bulrush Wm06

Schoenoplectus acutus

General Description Great bulrush marshes occur widely in subzones with warm and dry summers. Wave-exposed lake embayments with significant water movements, and grassland potholes with occasional substrate exposure (conditions that provide abundant aeration and limit organic accumulations), are the most common locations for this Site Association. Plant diversity is low; typically, Schoenoplec- tus acutus is the only species with significant cover. Bulrush marshes are usually adjacent to open water in wetland mosaics and can sometimes be found in complex with the Wm05. Floodwaters to 1.5 m depth in the spring are typical, with significant growing-season drawdown occurring in potholes. Great bulrush is tolerant of alkali soils and often dominates in brackish potholes. Soils are mostly Gleysols and Humic Gleysols, though Terric Humisols occasionally occur.

Characteristic Vegetation Wetland Edatopic Grid

Tree layer (0 - 0 - 0) Soil Nutrient Regime Shrub layer (0 - 0 - 5) ABCD E F Herb layer (10 - 70 - 100) Schoenoplectus acutus VM Moss layer (0 - 0 - 60) W Comments AkVA N MA SA The Wm06 includes marshes dominated by VW S. tabernaemontani (soft-stemmed bulrush). On wave-exposed lake shorelines or where Soil Moisture Regime pH sites are more brackish,S. acutus is more frequent,while in protected waters and St Sl Mo Dy VD potholes with mucky substrates, Hydrodynamic Index S. tabernaemontani is typical. Site conditions for Wm05 and Wm06 over- lap.S. acutus dominates on sites with alkaline mineral soils,greater wave exposure,or pronounced surface drying.Where marshes are heavily grazed by Muskrat,S. acutus is often favoured over Typha latifolia because it stores nutrients in the root mass and recovers more rapidly from grazing.

Chapter 5.3 Marshes 113 Wm07 Baltic rush

Juncus balticus

General Description Baltic rush saline meadows/marshes are common in the Chilcotin Plateau and uncommon in the dry climates of the Southern Interior and Southern Interior Mountains. The Wm07 occurs in alkaline or saline potholes, primarily closed basins, where there is early-season inundation followed by gradual watertable drop to below the surface. Juncus balticus is always dominant on Wm07 sites. Other saline-tolerant species such as Carex praegracilis, Potentilla anserina, and Puccinellia nuttalliana may occur, especially on drier sites. Soils are ﬁne textured, and poorly to imperfectly drained, with up to 10 cm of surface organic accumulation. The upper horizons remain wet throughout most of the growing season. When these sites dry out, a salt or alkali crust is often evident.

Characteristic Vegetation Wetland Edatopic Grid

Tree layer (0 - 0 - 0) Soil Nutrient Regime Shrub layer (0 - 0 - 2) ABCD E F Herb layer (15 - 70 - 100) Hordeum jubatum,Juncus balticus, VM Potentilla anserina Moss layer (0 - 5 - 20) W AM ANAkVA N MA SA Comments VW The Wm07,along with other communities of saline or alkaline soils conditions,requires Soil Moisture Regime pH site conditions that concentrate salts.These conditions are found in closed basins of semi- St Sl Mo Dy VD Hydrodynamic Index arid climates,where high evaporation rates and limited freshwater inflow lead to salt concentration.These same conditions also result in variable watertables within and between years,reflecting a changing balance of inflows and evaporation.Under these variable conditions the optimum environment for Site Associations changes location within the basin between years.Juncus balticus occupies those zones where flooding is shallow but soils do not completely dry out in the summer.Wm07 can form extensive stands in seasonally flooded depressions or as peripheral communities in the drawdown zone around permanent ponds and Wm06 marshes.Drier sites are Gs03 or Gs02.

114 Wetlands of British Columbia: A Field Guide to Identiﬁcation Sitka sedge – Hemlock-parsley Wm50

Carex sitchensis – Oenanthe sarmentosa

General Description Sitka sedge – Hemlock-parsley marshes are common in the Georgia Depression and Coast and Mountains at low elevations in basins, and along slow-moving streams, ponds, and lakeshores. A monoculture of Carex sitchensis occurs on many sites, but occasionally other species are also prominent. Sites with flowing floodwaters often have abundant Oenanthe sarmentosa or Glyceria elata. Drier sites have a mix of forbs such as Veronica scutellata, Hypericum anagalloides, and Galium trifidum. The Wm50 tolerates variable hydrology and disturbance and occurs on mineral substrates or shallow (occasionally deep) peat veneers.

Characteristic Vegetation Wetland Edatopic Grid

Tree layer (0 - 0 - 0) Soil Nutrient Regime Shrub layer (0 - 1 - 4) ABCD E F Herb layer (70 - 80 - 100) Carex sitchensis,Galium triﬁdum, VM Oenanthe sarmentosa Moss layer (0 - 0.1 - 0.5) W AM ANAkVA N MA SA Comments VW The Wm50 is the coastal equivalent of the Wm01 of the Interior. Soil Moisture Regime pH Ceska (1978) describes several variants of St Sl Mo Dy VD the Wm50,including Carex sitchensis with Hydrodynamic Index C. obnupta,Cicuta douglasii,and Aster subspicatus occurring adjacent to alder forests, with Deschampsia cespitosa,Gentiana sceptrum,and Hypericum anagalloides on drier sites,or with Glyceria elata and Calama- grostis canadensis in shaded areas with waterﬂow.

Chapter 5.3 Marshes 115 Wm51 Three-way sedge

Dulichium arundinaceum

General Description Three-way sedge marshes/fens are uncommon and often of limited extent on the south Coast and rare in wet regions of the Southern Interior Mountains at elevations below 600 m. The Wm51 occurs along the protected margins of shallow lakes or sluggish streams on mucky substrates. Dulichium arundinaceum is always dominant. There is often a minor component of emergent sedges or rushes. Other species that grow well on flooded, degrading peaty soils, such as Nuphar lutea, Menyanthes trifoliata, and Comarum palustre occur on some sites. The Wm51 prefers permanently flooded conditions on degrading peat or soft muck (mix of fine mineral material and organics).

Characteristic Vegetation Wetland Edatopic Grid

Tree layer (0 - 0 - 0) Soil Nutrient Regime Shrub layer (0 - 2 - 5) ABCD E F Herb layer (40 - 80 - 100) Carex sitchensis,Dulichium arundinaceum, VM Nuphar lutea Moss layer (0 - 5 - 10) W AM ANAkVA N MA SA Comments VW The Wm51 often occurs at the interface between peatland ecosystems such as the Soil Moisture Regime pH Wf52 and shallow-water ecosystems dominated by pond-lily or water shield. St Sl Mo Dy VD Hydrodynamic Index

116 Wetlands of British Columbia: A Field Guide to Identiﬁcation Additional Marsh Site Associations

This section briefly describes some uncommon Marsh Site Associations that have been sampled in British Columbia. Sharp bulrush Schoenoplectus pungens The Sharp bulrush Marsh Site Association is uncommon and restricted to the warm, dry subzones of the Southern Interior (BG, PP, and warmer IDF) at low elevations. These marshes are generally small and occur most commonly around small alkaline potholes or seepages where water levels are shallow and relatively constant. Rooting substrates are fine-textured mineral soil and can be saline/alkaline. Sites are always dominated by Schoenoplectus pungens with a scattering of other alkali-tolerant species such as Eleocharis palustris, Hordeum jubatum, Juncus balticus, and Triglochin maritima. Sharp bulrush marshes can occur adjacent to Wm06, Wm04, or Gs01 communities. Common reed Phragmites australis The Common reed Site Association is very uncommon and restricted to regions with warm summers such as the PP, BG, and hot IDF subzones of the Southern Interior and Southern Interior Mountains. Phragmites australis is a native rhizomatous grass that is more common in eastern Canada. It forms monocultures of tall stems (to 3 m) in shallow water of lakes, ponds, and slow-moving streams on mineral soils. Extreme watertable fluctuations are common, with deep flooding in spring to watertables well below the surface by the end of the growing season. Adjacent sites with more permanent soil saturation are occupied by Wm05 or Wm06. Inflated sedge Carex exsiccata Inflated sedge marshes occur widely but rarely throughout the southern two-thirds of the province at low to montane elevations. Carex exsiccata stands occur in conditions similar to those of C. utriculata. Sites have prolonged shallow flooding of mineral soils. Inflated sedge marshes have been sampled adjacent to small lakes, streams, and potholes. Northern mannagrass Glyceria borealis The Northern mannagrass Marsh Site Association is uncommon throughout the Interior at lower elevations. These marshes are often small and occur in shallow standing water at the margins of lakes, ponds, and slow-moving streams. Bottom substrates may be fine-textured mineral soils or well humified organic deposits. Glyceria borealis is the dominant species and may form a virtual monoculture. Such communities can be adjacent to Wm01 marshes, which occupy drier sites. (Described by Steen and Roberts 1988.)

Chapter 5.3 Marshes 117 Additional Marsh Site Associations

Reed canarygrass Phalaris arundinacea Reed canarygrass communities are common throughout the southern two-thirds of the province in areas with warm and relatively dry summers. They represent a disclimax community that establishes or is seeded on cleared willow swamps and low-bench sites. Typical sites are the floodplains of low-gradient streams or lake flats that are flooded in the spring and have prolonged soil saturation. Soils are commonly Gleysolic and fine-textured. Phalaris arundinacea is strongly rhizotomous and produces a dense sod and full canopy that excludes most other species; most stands are monotypic. Seacoast bulrush Bolboschoenus maritimus The Seacoast bulrush Marsh Site Association is uncommon and restricted to the dry subzones of the Southern Interior at low to middle elevations. These marshes are generally small and occur most commonly around alkaline potholes. Soil reaction is generally > 8.5 pH and may also be saline. Soils are fine-textured and continually saturated. Bol- boschoenus maritimus usually forms moderately dense monocultures. The Wm06 or shallow-water ecosystems may occur in adjacent more deeply flooded habitats. Seacoast bulrush communities also occur in southern estuaries (see Chapter 5.6). Seaside arrow-grass Triglochin maritima The Seaside arrow-grass Marsh Site Association is uncommon and restricted to the dry subzones of the Central Interior at low elevations. These marshes are generally very small and occur most commonly around small saline potholes. Sites are seasonally inundated, though standing water can persist until late in the season. Soils are fine-textured Gleysols with thin, well-humified organic layers. Sites have very low species diversity; frequently only Triglochin maritima occurs. This ecosystem may represent a short-lived successional community that establishes on saline flats experiencing “improved” water regime. (Described by Steen and Roberts 1988.) Woolly sedge Carex lanuginosa The Woolly sedge Marsh Site Association is uncommon and restricted to the warm, dry subzones of the Central and Southern Interior (BG, PP, and warmer IDF) at low elevations. These marshes are generally small and occur in small freshwater, grassland-surrounded potholes where water levels are shallow and relatively constant. They are also found in small patches within some larger Wm01 or Wm06 marshes. Standing

118 Wetlands of British Columbia: A Field Guide to Identiﬁcation Additional Marsh Site Associations

water is slightly alkaline; rooting substrates are ﬁne-textured mineral soil. Sites are always dominated by Carex lanuginosa but infrequently have other species, such as Drepanocladus aduncus, Myriophyllum verticillatum, Alopecurus aequalis, Carex atherodes, or C. utriculata, occurring with signiﬁcant cover. Wm03 marshes occur in similar hydrological conditions but are more common in the forested zones.

1 Typha latifolia, common cattail 2 Schoenoplectus acutus, great bulrush 3 Equisetum fluviatile, swamp horsetail

Chapter 5.3 Marshes 119 5.4 SWAMPS

1 Red alder - Skunk cabbage swamp, Vancouver Island (CWHxm1) 2 A Spruce – Horsetail swamp at Mount Savona near Kamloops (MSxk) 3 MacCalla’s willow – Beaked sedge swamp near Uncha Lake, Sub-Boreal Interior (SBSdk)

Chapter 5.4 Swamps 121 SWAMP WETLAND CLASS

Definition A swamp is a nutrient-rich wetland ecosystem where significant groundwater inflow, periodic surface aeration, and/or elevated microsites allows growth of large trees or tall shrubs under subhydric conditions. General Description Vegetation Table 5.4.2 lists common species of Swamp Site Associations described in this guide. Swamps are characterized by a high cover of tall shrubs and/or trees plus a well-developed herb layer. Richer swamps have a herb layer with a high component of ferns and forbs. Nutrient-medium sites have a sedge-dominated understorey. There are two distinct groups of swamps: one characterized by a tall- shrub physiognomy and the other forested. The former are often floristically related to fen ecosystems but distinguished by vigorous shrub growth. The moss layer is typically poorly developed because shade and abundant litterfall limit bryophyte establishment and growth. Forested swamps are transitional to uplands and often have a mix of terrestrial and wetland microhabitats. Elevated microsites under conifers are favourable for terrestrial species, and depressions support hydrophytes. Landscape Position and Distribution Swamps occur where there is an abundant flow of near-surface groundwater, and on slope breaks, peatland margins, inactive floodplain back-channels, back-levee depressions, lake margins, and gullies. Swamps are common throughout the province, but usually not extensive in most landscapes (Table 5.4.1). Most frequently, they occur as small components of larger wetland systems. Hydrology and Soils Swamps occur on sites with pronounced microtopography. Trees and shrubs root on microsites elevated above a high semipermanent watertable or where there is pronounced lateral groundwater flow and surface aeration in otherwise saturated soils (Figure 5.4.1).

122 Wetlands of British Columbia: A Field Guide to Identification Soils are usually distinctly gleyed. Soil Nutrient Regime A surface horizon of well-decom- ABCD E F posed woody peat smelling M strongly of hydrogen sul de is fi VM common. Occasionally, swamps occur on deeper peat deposits but W Swamps this is the exception; these are AkVA N MA SA either peatlands that have under- VW gone hydrological change or are Soil Moisture Regime pH sites fed by well-aerated water that provides enough oxygen for St Sl Mo Dy VD Hydrodynamic Index growth of larger plants. Swamps have abundant available nutrients  .. Position of swamps on the supplied by groundwater flow. edatopic grid. Other Comments Most of the forested Swamp Site Associations presented in this section are transitional to upland ecosystems and have been previously described in  field guides (e.g., Banner et al. 1993). The plant communities on these sites do not have an abundance of obligate hydrophytes and do not clearly distinguish sites that are on wetland soils from those on merely very moist upland soils at this ecotone. Therefore, such Site Associations include some productive upland sites and also non-productive sites. “Poor swamps,” those sites with trees, stagnant hydrology, relatively few miner0trophic indicators, and abundant bog-affiliated species, are included in the Bog Class. Conservation Issues Swamps are important habitats for wildlife. They have more vertical structure than other wetland classes and therefore support a more diverse avifauna. Species that use seral stands for nesting and feeding also use shrub swamps. Forested swamps have a characteristically open or patchy canopy that appears to be favoured by many bird and bat species. Several important bear foods, such as Lysichiton americanus and Equise- tum arvense, are common in swamps. Willow and other shrubs are important browse for ungulates. Livestock and wildlife use swamps for browse, shade, and cover, especially where they occur adjacent to open grassland or sources of drinking water. Excessive grazing will affect these sites, and rest rotation methods are recommended. In many areas where shrub swamps are cleared for

Chapter 5.4 Swamps 123 pasture, dense and persistent stands of reed canarygrass (Phalaris arundinacea) become established. Forested swamps are productive relative to other wetland ecosystems but are still marginal for timber production. The forested ecosystems described in this guide occur at the limits of operability. Some sites are dry enough and productive enough to be successfully regenerat- ed but these are in the minority. Sites with large trees on wetland soils are especially risky for timber harvest since watertables rise post-harvest, microtopography (which allows large tree growth) is easily destroyed, and brush competition is high. Operable and inoperable sites share essentially the same indicator species groups, however, and so are not separated at the Site Association level.

Spiraea douglasii, hardhack or pink spirea

124 Wetlands of British Columbia: A Field Guide to Identification xx x x xxx xx xx xx w w w w w w xxxxx BGPP BWBS SWB ESSF ICH IDF MS SBS CDF CWH SBPS MH Distribution of Swamp Site Associations by biogeoclimatic zone Mountain alder – Skunk cabbage – Lady fern alder – Skunk cabbage Lady Mountain sedge – Sitka alder – Pink spirea Mountain willow – BluejointBebb’s sedge willow – Beaked Drummond’s sedge – Beaked willow MacCalla’s sedge – Sitka willow Sitka moss horsetail – Leafy – Common Spruce valerian horsetail fir – Sitka – Common Subalpine – Skunk cabbage Peat-mossBlack spruce – Skunk cabbage – Spruce redcedar Western fir – Skunk cabbage – Subalpine Spruce sedge – Sitka Pink spirea willow – Skunk cabbage willow – Pacific Sitka x alder – Skunk cabbage Red fern – Skunk cabbage – Sword redcedar Western x hemlock – Skunk cabbageWestern – redcedar Western xx hemlock – Skunk cabbage – Mountain Yellow-cedar xx x xx x xx xx x x xx xx x xx xx xx xx xx x xxx xx x x xx x x x x x x x x x xx x x x = incidental; < 5% of wetlands xx = minor; 5–25% of wetlands xxx = major; >25% of wetlands w = wet subzones only x = very dry subzones only Ws01 Ws02 Ws03 Ws04 Ws05 Ws06 Ws07 Ws08 Ws09 Ws10 Ws11 Ws50 Ws51 Ws52 Ws53 Ws54 Ws55  .. 

Chapter 5.4 Swamps 125  .. Swamp Species Importance Table

Species Ws03 Ws04 Ws05 Ws02 Ws06 Ws07 Ws08 Ws01 Trees Picea X x x x x x xyzzz xyzzz xyz Picea mariana x Abies lasiocarpa x xy xyzzzz xy Tsuga heterophylla x Thuja plicata x Picea sitchensis Alnus rubra Acer macrophyllum Chamaecyparis nootkatensis Tsuga mertensiana x Abies amabilis Shrubs Salix bebbiana xyzzzz x x Salix drummondiana x xyzzz x x Salix maccalliana x x xyzzzz Alnus incana xyzz x xyzzz xyz xyz xyzzzz Lonicera involucrata xyz xy xy xy xy xyzz xy xyz Spiraea douglasii x xy xyzz xy xyzz Cornus stolonifera xyz x xy xy xy Vaccinium alaskaense/ovalifolium xy Salix sitchensis xy xyzzzz Salix lucida xx Rubus spectabilis x Sambucus racemosa x Gaultheria shallon Ribes bracteosum Elliottia pyroliﬂorus Herbs Calamagrostis canadensis xyzzz xyz xyzz xyzz xyz xy xy xyz and Carex aquatilis/sitchensis x xyzz xyzz xyzzzz xyzzz xy Dwarf Carex utriculata xyz xyzz xyzzz xy xyz Shrubs Gymnocarpium dryopteris xxxxyxyxy Valeriana sitchensis xyzz Scirpus microcarpus x xyz Equisetum arvense xyzz x x xy xyzz xyzzzz xyzzz xyz Lysichiton americanus x xy xyzzzz Athyrium ﬁlix-femina x xy x x xyzzz Tiarella trifoliata xy xy x Streptopus lanceolatus xx Maianthemum dilatatum Oenanthe sarmentosa Polystichum munitum Equisetum telmateia Blechnum spicant Veratrum viride xxyx Fauria crista-galli Mosses Drepanocladus spp. x xyzz x x and Mnium spp. xyz xy xyz xyz xy xyzzz xyzzz xyzz Lichens Aulacomnium palustre x xy xyz x xyzz xyzz x Sphagnum spp. xxx x x Hylocomium splendens x xyzz Pleurozium schreberi x xyz xy x Eurhynchium praelongum Rhytidiadelphus loreus

126 Wetlands of British Columbia: A Field Guide to Identification Ws09 Ws10 Ws11 Ws50 Ws51 Ws52 Ws53 Ws54 Ws55 Common Name xyz xyzz xyzz spruce xyzz x x black spruce xyzzz xyzz xyzzzz subalpine fir xy xyzzz x xyzz xyzzzz xyzzzz western hemlock x xyzzzz x xyz xyzzzz xyzzz xy western redcedar xy x xyz xyzz x Sitka spruce x xy xyzzzz xyzzz xy x red alder xyz xyzz bigleaf maple xy xyzzzz yellow-cedar x xyzzzz mountain hemlock xyz xyzzz amabilis fir x Bebb’s willow x Drummond’s willow MacCalla’s willow xyzz xy xyzzz x mountain alder xy xy xyzz x xyz x x x black twinberry xyzz x xyz xyzzzz xyz x x pink spirea x xy xy x xyz xy xy x red-osier dogwood xyz xyzz xyz x xyzzz xyzzzz Alaska/oval-leaved blueberry x xyzzzz Sitka willow x xyzzzz Pacific willow x x xyzz xyzzzz xyzzz xyzz xyz salmonberry x x xyzzz xyzz x red elderberry x x xy xyzz x salal x xyzz xy x stink currant xyz copperbush xyz xy xy x x x x bluejoint x xyzz x x water/Sitka sedge xy beaked sedge xy xyzz xyzz x x xy xy oak fern Sitka valerian xx xyzxx small-flowered bulrush xyzz xyzz xyzz x xyz x x x common horsetail xyzzzz xyzzz xyzzzz xy xyzzzz xyzzzz xyzzz xyzzz xyzzz skunk cabbage xyz xyzzz xyzzz x xyzz xyzzz xyzzz xyzz xyz lady fern x xyz xyzz x xyz xyz xyz xy foamflower xy xyz xyz x x xy xy rosy twistedstalk x xyzz xyzz xy xy false lily-of-the-valley x xyzz xy x x Pacific water-parsley xyz xyzz x x sword fern x xy xyzz x giant horsetail x x x xyzz xyz deer fern x x x x xyz Indian hellebore x xyz deer-cabbage x hook-mosses xyz xyzzz xyzzz x xyz xyzz xyzz xyzz xyzz leafy mosses xy x xy x glow moss xyzzz xyzz xyz xy x x xyz xyzz peat-mosses x xyz x x x xyzz xyz step moss xyzz xyz xy red-stemmed feather-moss xy xyzzz xyzzz xy x beak moss x x x x xyzz xyzz lanky moss

Chapter 5.4 Swamps 127 Ws01 Mountain alder – Skunk cabbage – Lady fern

Alnus incana – Lysichiton americanus – Athyrium ﬁlix-femina

General Description Mountain alder – Skunk cabbage – Lady fern swamps are common in wet regions of the Sub-Boreal Interior and South- ern Interior Mountains, particularly in areas underlain by glaciolacustrine deposits. The Ws01 frequently occurs in wet gullies or along small creeks where there is continuous seepage near the surface and poor drainage. It also occurs in the lagg of peatlands, where seepage from upslope enriches peat deposits. Alnus incana dominates these sites, which have a lush and diverse understorey where Athyrium filix-femina and Lysichiton americanus are prominent. Scattered spruce is common. The moss layer is often sparse because of shading and high rates of litterfall. Soils are usually poorly drained, fine-textured mineral deposits with a veneer of well-humified woody peat. Occasionally this unit will occur on deeper peat deposits.

Characteristic Vegetation Wetland Edatopic Grid Tree layer (0 - 5 - 8) Soil Nutrient Regime Picea X ABCD E F Shrub layer (20 - 53 - 99) Alnus incana,Lonicera involucrata,Picea X, VM Spiraea douglasii Herb layer (35 - 68 - 95) W Athyrium ﬁlix-femina,Calamagrostis AkVA N MA SA canadensis,Equisetum arvense, Lysichiton americanus VW Moss layer (0 - 24 - 87) Mnium spp. Soil Moisture Regime pH

St Sl Mo Dy VD Comments Hydrodynamic Index Several other mountain alder–dominated Site Associations occur.The Fl01 and Fl02 occur on well-drained soils adjacent to streams and rivers; these sites lack skunk cabbage.On wetter sites,Carex sitchensis is dominant in the understorey and described by the Ws02. The Ws01 often fully occupies small depressions and gullies in upland forest.It also occurs between sedge fens and upland forest.

128 Wetlands of British Columbia: A Field Guide to Identiﬁcation Mountain alder – Pink spirea – Sitka sedge Ws02

Alnus incana – Spiraea douglasii – Carex sitchensis

General Description The Mountain alder – Pink spirea – Sitka sedge Swamp Site Association is common in wet climates of the Sub-Boreal In- terior, Southern Interior Mountains, and interior transition areas of the Coast and Mountains. The Ws02 occurs on beaver-flooded flats of small creeks, peripheral zones of wetlands, and lakeshores, where there is early season flooding, continuous seepage near the surface, and poor drainage. Alnus incana forms an open to sparse canopy. Spiraea douglasii can be scattered or prominent. Carex aquatilis or C. sitchensis is usually the dominant species of the herb layer, but Scirpus microcarpus dominates on some sites. Significant cover of Calamagrostis canadensis is common. Soils are derived from fluvial or lacustrine material and often have a veneer or blanket of sedge peat. Organic horizons have silty or sandy lenses throughout, indicating periodic significant flood events.

Characteristic Vegetation Wetland Edatopic Grid

Tree layer (0 - 0 - 2) Soil Nutrient Regime Shrub layer (12 - 44 - 99) ABCD E F Alnus incana,Spiraea douglasii Herb layer (20 - 72 - 100) VM Calamagrostis canadensis,Carex aquatilis/sitchensis,Comarum W palustre,Scirpus microcarpus AkVA N MA SA Moss layer (0 - 10 - 85) Mnium spp. VW

Soil Moisture Regime pH Comments

St Sl Mo Dy VD The Ws02 is similar to Willow – Sedge Site Hydrodynamic Index Associations but occurs on sites with more dynamic water ﬂow; willow-dominated sites (Ws04–06) tend to be more stagnant.

Chapter 5.4 Swamps 129 Ws03 Bebb’s willow – Bluejoint

Salix bebbiana – Calamagrostis canadensis

General Description Bebb’s willow – Bluejoint swamps are uncommon but widespread throughout the drier climates of the Interior at elevations below 1200 m. They occur on lake flats, pond margins, fluvial terraces, seasonal creeks, and palustrine basins where early-season shallow standing water draws down to very moist conditions by late growing season. Salix bebbiana forms an open canopy, often with a significant component of Alnus incana. Scat- tered spruce trees can occur. Various other shrub species are common in the understorey. Calama- grostis canadensis is a constant dominant but usually occurs only on raised microsites. Equisetum arvense and other horsetails can also be prominent. A diversity of other forbs with low cover is typical. Sites often have distinct mounds created by fallen trees, interspersed with sparsely vegetated pools of water; however, some stands are drier and have a more continuous herbaceous understorey. Soils are fine-textured Gleysols, often with veneers of woody peat.

Characteristic Vegetation Wetland Edatopic Grid

Tree layer (0 - 2 - 5) Soil Nutrient Regime Shrub layer (20 - 46 - 99) ABCD E F Alnus incana,Cornus stolonifera, Lonicera involucrata,Salix bebbiana VM Herb layer (6 - 56 - 90) Calamagrostis canadensis,Carex utriculata, W Equisetum arvense AkVA N MA SA Moss layer (0 - 19 - 75) Mnium spp. VW Soil Moisture Regime Comments pH

The Ws03 has soils and hydrology characteris- St Sl Mo Dy VD tic of wetland ecosystems but typically has Hydrodynamic Index low cover of obligate hydrophytes.

130 Wetlands of British Columbia: A Field Guide to Identiﬁcation Drummond’s willow – Beaked sedge Ws04

Salix drummondiana – Carex utriculata

General Description Drummond’s willow – Beaked sedge swamps/fens are common in the Central and Sub-Boreal Interior in back-levee depressions of low-gradient creeks or channel margins in peatland streams. Ws04 sites can be deeply flooded during the spring freshet and after drawdown maintain a high watertable due to fine-textured soils or low- lying position relative to the watertable. Salix drummondiana dominates these sites, with other shrubs such as Lonicera involucrata and Spiraea douglasii common in the low- shrub layer. The herb layer is moderately well developed and predominantly Carex aquatilis and C. utriculata. Sedge peat veneers or blankets over fine- to medium-textured fluvial or lacustrine materials are typical. Flooding can result in buried organic layers, peat and mineral mixing, or reduced surface organic accumulation.

Characteristic Vegetation Wetland Edatopic Grid

Tree layer (0 - .5 - 10) Soil Nutrient Regime Shrub layer (10 - 52 - 100) ABCD E F Lonicera involucrata,Salix drummondiana, Spiraea douglasii VM Herb layer (2 - 53 - 90) Calamagrostis canadensis,Carex W aquatilis/sitchensis,C. utriculata AkVA N MA SA Moss layer (0 - 14 - 80) Mnium spp. VW Soil Moisture Regime Comments pH

In wetter climates of the Interior,the Ws04 is St Sl Mo Dy VD Hydrodynamic Index replaced by the Ws06. The Ws04 occurs along open water channels adjacent to Wf01 and Wm01 units.It also occurs in low sites along sluggish streams adjacent to the Fl05. Drummond’s willow is well adapted to ﬂuvial sites; twigs and branches have brittle bases that readily break during ﬂood events.These whips will readily root in mineral soils.

Chapter 5.4 Swamps 131 Ws05 MacCalla’s willow – Beaked sedge

Salix maccalliana – Carex utriculata

General Description MacCalla’s willow – Beaked sedge swamps/fens occur in scattered locations in drier climates of the Central and Sub-Boreal Interior in basins, hollows, and streamside areas that are shallowly flooded in the early season by slowly flowing waters. Sites often have complex microtopography with tall willows rooting on elevated hummocks, and with depressions with standing water. Tall Salix maccalliana dominates these sites but a diversity of other shrubs is common. Carex utriculata or C. aquatilis are usually dominant in the understorey but because of the pronounced microtopography a diversity of species often occurs. The moss layer is often moderately developed. Soils are variable, ranging from deep mesic peat to thin layers of humic muck. Peat accumulations from 20 to 400 cm with well-humified surface tiers are typical.

Characteristic Vegetation Wetland Edatopic Grid

Tree layer (0 - .1 - 2) Soil Nutrient Regime Shrub layer (25 - 60 - 85) ABCD E F Betula nana,Salix glauca,S. maccalliana Herb layer (13 - 54 - 95) VM Calamagrostis canadensis, Carex aquatilis, C. utriculata W Moss layer (0 - 40 - 100) AkVA N MA SA Aulocomnium palustre,Drepanocladus spp., Mnium spp. VW

Soil Moisture Regime pH Comments

Pronounced lateral water flow in Ws05 sites St Sl Mo Dy VD Hydrodynamic Index allow the robust growth of Salix maccalliana on peaty soils. Sites occasionally occur on deep deposits of sedge peat with a humic surface tier,suggesting that these sites have developed on hydrologically modified fens. Sites with more active flooding are occupied by the Ws04.

132 Wetlands of British Columbia: A Field Guide to Identiﬁcation Sitka willow – Sitka sedge Ws06

Salix sitchensis – Carex sitchensis

General Description Sitka willow – Sitka sedge swamps are uncommon at low elevations in the Coast and Mountains, Nass Basin, and wet subzones of the Southern Interior Mountains and Sub-Boreal Interior. These sites are usually associated with fluvial systems or linked basins and experience prolonged saturation and brief early-season flooding. Salix sitchensis dominates Ws06 sites. The herb layer is primarily Carex sitchensis and Equisetum arvense. Other large sedges and forbs are also common. On some sites, particularly those under shade, Scirpus microcarpus replaces C. sitchensis as the site dominant. The moss layer is poorly developed. Gleysols derived from fluvial materials are the most common soil type. On some sites, sedge peat is layered in fluvial deposits.

Characteristic Vegetation Wetland Edatopic Grid

Tree layer (0 - .2 - 2) Soil Nutrient Regime Shrub layer (15 - 50 - 90) ABCD E F Alnus incana,Salix sitchensis Herb layer (30 - 74 - 99) VM Calamagrostis canadensis,Carex sitchensis, C.utriculata,Equisetum arvense, W Scirpus microcarpus AkVA N MA SA Moss layer (2 - 8 - 35) Mnium spp. VW Soil Moisture Regime Comments pH

Adjacent communities are often Wm01 or St Sl Mo Dy VD Hydrodynamic Index Wm02 marshes or low bench flood communities.This Site Association is similar to the Ws04 and Ws02; the former occurs in drier subzones and the latter on more active floodplain sites. Sitka willow is well adapted to fluvial sites; twigs and branches have brittle bases that readily break during flood events.These whips will readily root in mineral soils.

Chapter 5.4 Swamps 133 Ws07 Spruce – Common horsetail – Leafy moss

Picea X – Equisetum arvense – Mnium

General Description The Spruce – Common horsetail – Leafy moss Swamp Site As- sociation is common in the Northen Boreal Mountains and Cen- tral and Sub-Boreal Interior from low to subalpine elevations. It occurs on lower and toe slopes and margins of wetlands, where there is significant flow of mineral-rich groundwater. These can be moderately productive sites with spruce to 25 m tall rooting on elevated mounds. The shrub layer may be well developed or sparse with Lonicera involucrata the most prominent species. Equise- tum arvense is always abundant but a diversity of other upland and wetland species is common. The moss layer can be diverse, with leafy mosses (Mnium spp.) and Aula- comnium palustre usually prominent in depressions and Pleurozium schreberi and other feathermosses on raised mounds. Soils most often have a thin, dark, well-humified, woody peat veneer over fine-textured mineral soils but occasionally deeper peat deposits are encountered. Characteristic Vegetation Wetland Edatopic Grid Tree layer (10 - 25 - 50) Picea X Soil Nutrient Regime Shrub layer (25 - 30 - 70) ABCD E F Alnus incana,Lonicera VM involucrata,Picea X Herb layer (6 - 70 - 90) Equisetum arvense W Moss layer (5 - 70 - 99) AkVA N MA SA Aulacomnium palustre,Hylocomium splendens,Mnium spp.,Pleurozium VW

schreberi Soil Moisture Regime pH

Comments St Sl Mo Dy VD Hydrodynamic Index Spruce – Horsetail units have been described for many interior biogeoclimatic subzones in regional ﬁeld guides (see Appendix 4).Most of these Site Series include sites with freely drained soils supporting productive forests,as well as stands with wetland soils and poor tree productivity.Plant community composition does not clearly reﬂect these separate conditions because most species are facultative wetland indicators,and pronounced microtopography allows upland species to occur on wetland sites.The Ws07 therefore includes wetland and non-wetland sites.Wetland sites will have poor tree productivity and hydric soils. A similar site with more stagnant hydrology and greater peat development is the Wb08.Ws08 ecosystems are generally similar to the Ws07 but occur at high elevations and have abundant subalpine indicators.

134 Wetlands of British Columbia: A Field Guide to Identiﬁcation Subalpine ﬁr – Sitka valerian – Common horsetail Ws08

Abies lasiocarpa – Valeriana sitchensis – Equisetum arvense

General Description Subalpine fir – Sitka valerian – Common horsetail swamps are common at elevations above 1100 m throughout the Interior. The Ws08 occurs on lower and toe slopes and margins of wetlands, where there is significant flow of mineral-rich groundwater. The canopy is open and patchy with groups of interior spruce and subalpine fir separated by forb-rich openings. The shrub layer may be well developed or sparse. The herb layer is generally well developed; an abundance of Equisetum arvense and subalpine forbs is typical. Leafy mosses (Mnium spp.) and Aula- comnium palustre are usually prominent in depressions and Barbilophozia and feathermoss species are prominent on raised mounds. Gleysolic soils have a thin, dark, well-humified, woody peat veneer over fine-textured mineral soil, but occasionally deeper peat deposits are encountered.

Characteristic Vegetation Wetland Edatopic Grid Tree layer (6 - 40 - 80) Abies lasiocarpa,Picea X Soil Nutrient Regime Shrub layer (10 - 40 - 80) ABCD E F Abies lasiocarpa,Picea X,Vaccinium VM membranaceum Herb layer (25 - 70 - 99) W Equisetum arvense,Rubus pedatus, Senecio triangularis,Streptopus AkVA N MA SA amplexifolius,Valeriana sitchensis Moss layer (5 - 70 - 90) VW Aulacomnium palustre,Barbilophozia Soil Moisture Regime pH lycopodioides,Brachythecium spp., Mnium spp.,Pleurozium schreberi St Sl Mo Dy VD Hydrodynamic Index Comments Subalpine fir – Horsetail units have been described for many ESSF subzones in regional field guides (see Appendix 4).Many subzones have two Subalpine fir – Horsetail site series described; one for poor sites and one for rich sites.Rich sites have abundant ferns,only occur at lower elevations of the ESSF,and more closely resemble Spruce – Horsetail sites.The Ws08 does not include these sites. The Ws08 is the high-elevation equivalent of the Ws07.Tree growth is disproportionately slow at higher elevations because of cold conditions in wet soils and persistent snowpack.With increasing elevation,open patches with abundant subalpine forbs become larger and trees become more restricted to elevated sites.

Chapter 5.4 Swamps 135 Ws09 Black spruce – Skunk cabbage – Peat-moss

Picea mariana – Lysichiton americanus – Sphagnum

General Description Black spruce – Skunk cabbage – Peat-moss poor swamps/bogs are uncommon in the “rainforest” climate areas of the Central Interior (wet SBS and northern ICH), in palustrine basins and back-levee depressions with high watertables. These sites are strongly mounded, with conifers on elevated microsites and standing water in between. Canopy composition is diverse with Picea mariana, Abies lasiocarpa, Pinus contorta, and Tsuga heterophylla often all present. Pronounced microtopography can result in diverse, well-developed shrub, herb, and moss layers. Lysichiton americanus is always present in damp hollows and Sphag- num species dominate the moss layer. Organic veneers of dark woody humic or mesic peat over ﬁne-textured lacustrine material are typical. However, peat depths vary from 20 to 150 cm on different sites. Soils are Terric Humisols/Mesisols or Humic Gleysols with peaty veneers.

Characteristic Vegetation Wetland Edatopic Grid

Tree layer (0 - 18 - 35) Soil Nutrient Regime Abies lasiocarpa,Picea X,P.mariana ABCD E F Shrub layer (25 - 49 - 70) Abies lasiocarpa,Alnus incana,Ledum VM groenlandicum,Picea mariana,Spiraea douglasii,Vaccinium ovalifolium W Herb layer (6 - 56 - 90) AkVA N MA SA Athyrium ﬁlix-femina,Calamagrostis canadensis,Carex disperma,Equisetum VW arvense,Lysichiton americanus Moss layer (2 - 46 - 98) Soil Moisture Regime pH Mnium spp.,Pleurozium schreberi,Ptilium St Sl Mo Dy VD crista-castrensis,Sphagnum Group I, Hydrodynamic Index Sphagnum Group II

Comments Ws09 swamps occur in regions where Ws10 and Ws11 swamps also occur; however,this unit occurs on wetter sites and usually in locations with cold-air ponding.These sites are transitional to bogs but have some rich site indicators and have swamp-like soils and tree growth potential.The Wb08 is the equivalent Site Association of drier climates. The Ws09 has not been previously described in regional BEC ﬁeld guides,where it is lumped with the richer and more productive skunk cabbage forests of the ICH (Ws10) or SBS (Ws11).

136 Wetlands of British Columbia: A Field Guide to Identiﬁcation Western redcedar – Spruce – Skunk cabbage Ws10

Thuja plicata – Picea X – Lysichiton americanus

General Description Western redcedar – Spruce – Skunk cabbage swamps are uncommon in ICH zones of the Central Interior and Southern Interior Mountains. They occur on toe slopes, peatland margins, and low-lying areas in floodplains. Canopy composition is typically a mix of Picea X, Thuja plicata, and Tsuga heterophylla, with Abies lasiocarpa occurring in cold-air ponding sites. Pronounced microtopography can result in diverse, well-developed shrub, herb, and moss layers. Lysichiton americanus is always present in damp hollows, and rich-site indicators such as Gymnocarpium dryopteris, Athyrium filix-femina, and Equisetum arvense are abundant. Leafy mosses are prominent in a diverse moss layer. Organic veneers of dark woody humic or mesic peat over fine-textured lacustrine material are typical. However, peat depths are variable, ranging from 10 to 200 cm. Soils are Gleysols/Humic Gleysols with peaty humus forms or Terric Humisols/Mesisols.

Characteristic Vegetation Wetland Edatopic Grid

Tree layer (0 - 52 - 99) Soil Nutrient Regime Abies lasiocarpa,Picea X,Thuja plicata, ABCD E F Tsuga heterophylla Shrub layer (5 - 55 - 99) VM Oplopanax horridus,Thuja plicata,Tsuga heterophylla,Vaccinium ovalifolium W Herb layer (6 - 59 - 90) AkVA N MA SA Athyrium ﬁlix-femina,Cornus canadensis, Equisetum arvense,Gymnocarpium dry- VW opteris,Lysichiton americanus,Streptopus lanceolatus,Tiarella trifoliata Soil Moisture Regime pH Moss layer (5 - 63 - 98) Mnium spp.,Pleu- St Sl Mo Dy VD rozium schreberi,Ptilium crista-castrensis, Hydrodynamic Index Rhytidiadelphus triquetrus,Sphagnum Group II

Comments The Ws10 is the most common forested skunk cabbage Site Association in the Interior.Soils indicate a wetland environment but plant community composition does not clearly reﬂect this because pronounced microtopography allows upland species to occur.Most species in Ws10 sites are facultative wetland indicators. The Ws10 describes rich,wet skunk cabbage forests of the ICH; similar forests in the SBS are described by the Ws11.The Ws09 is similar but has more stagnant hydrology,greater peat development,and few rich-site indicators. The Ws10 includes several BEC Sites Series (see Appendix 4).

Chapter 5.4 Swamps 137 Ws11 Spruce – Subalpine ﬁr – Skunk cabbage

Picea X – Abies lasiocarpa – Lysichiton americanus

General Description Spruce – Subalpine fir – Skunk cabbage swamps are uncommon in the wet SBS subzones of the Sub-Boreal Interior on toe slopes, peatland margins, and low-lying areas in floodplains. Canopy composition is typically a mix of Picea X and Abies lasiocarpa. Pronounced microtopography can result in diverse, well-developed shrub, herb, and moss layers. Lysichiton americanus is always present in damp hollows, and rich- site indicators such as Gymnocarpium dryopteris, Athyrium filix-femina, and Equise- tum arvense are abundant. Organic veneers of dark woody humic or mesic peat over fine-textured lacustrine material are typical. However, peat depths are variable from 5 to 80 cm on different sites. Soils are Gleysols/Humic Gleysols with peaty humus forms or Terric Humisols.

Characteristic Vegetation Wetland Edatopic Grid

Tree layer (0 - 15 - 25) Soil Nutrient Regime Abies lasiocarpa,Picea X ABCD E F Shrub layer (30 - 41 - 60) Abies lasiocarpa,Alnus incana,Lonicera VM involucrata,Oplopanax horridus,Picea X, Spiraea douglasii,Vaccinium ovalifolium W Herb layer (40 - 59 - 95) AkVA N MA SA Athyrium ﬁlix-femina,Cornus canadensis, Drypopteris expansa,Equisetum arvense, VW Gymnocarpium dryopteris,Lysichiton americanus,Streptopus lanceolatus, Soil Moisture Regime pH Tiarella trifoliata St Sl Mo Dy VD Moss layer (2 - 23 - 40) Hydrodynamic Index Brachythecium spp.,Mnium spp., Sphagnum spp.

Comments The Ws11 is similar to the Ws10 but occurs in climatic areas where redcedar and hemlock are not present.Soils indicate a wetland environment but plant community composition does not clearly reﬂect this because pronounced microtopography allows upland species to occur.Most species in Ws11 sites are facultative wetland indicators. The Ws11 describes rich,wet,skunk cabbage forests of the SBS; similar forests in the ICH are described by the Ws10.The Ws09 is similar but has more stagnant hydrology,greater peat development,and few rich-site indicators. The Ws11 includes Site Series SBSvk/10.

138 Wetlands of British Columbia: A Field Guide to Identiﬁcation Pink spirea – Sitka sedge Ws50

Spiraea douglasii – Carex sitchensis

General Description Pink spirea – Sitka sedge swamps are common at low elevations of the Georgia Depression in basins, gullies, and margins of waterbodies and peatlands. These sites experience prolonged saturation and brief early-season ﬂooding. Species diversity is low in this Site Association. Spiraea douglasii always dominates Ws50 sites; few other shrub species occur. The sedge-dominated understorey is sparse or well developed. Few species other than Carex sitchensis are common. The moss layer is often minimal but Aulacomnium palustre or Sphagnum spp. occur with high abundance on some sites. Humisols and Gleysols are the most common soil types.

Characteristic Vegetation Tree layer (0 - 0 - 2) Shrub layer (15 - 70 - 99) Spiraea douglasii Herb layer (2 - 35 - 85) Carex sitchensis Moss layer (0 - 34 - 90) Aulacomnium palustre,Sphagnum Group I

Comments Wetland Edatopic Grid

The Ws50 Site Association is common as a Soil Nutrient Regime component of many peatlands along the ABCD E F southern Coast.It can be the dominant Site Association in small basins or surrounding VM Wm50 marshes.S. douglasii increases with disturbance and many spirea thickets actual- W ly represent disturbance communities that AkVA N MA SA have developed after hydrological change. Understoreys in these successional communi- VW ties vary from completely absent to bog-like. Soil Moisture Regime pH Shrub thickets dominated by Myrica gale with sedge are common in the region where St Sl Mo Dy VD the Ws50 occurs.These communities are usu- Hydrodynamic Index ally on peat and are described by the Wf52 Site Association.

Chapter 5.4 Swamps 139 Ws51 Sitka willow – Paciﬁc willow – Skunk cabbage

Salix sitchensis – Salix lucida – Lysichiton americanus

General Description Sitka willow – Pacific willow – Skunk cabbage swamps occur sporadically at low elevations throughout the Coast and Moun- tains, Georgia Depression, and coastal transition areas of the Interior at peatland margins and in floodplain depressions. Salix sitchensis and S. lucida often co-dominate a closed canopy of tall shrubs and low trees. The understorey is lush and dominated by Lysichiton americanus and Athyrium filix-femina. Wetter microsites have Scirpus microcarpus, Oenanthe sarmentosa, and Equisetum spp. The moss layer is typically sparse. Soils are mostly Gleysols with dark peat veneers. In palustrine locations, deeper humic organic deposits are common.

Characteristic Vegetation Wetland Edatopic Grid

Tree Layer (0 - 10 - 85) Soil Nutrient Regime Alnus rubra,Salix lucida,S.sitchensis ABCD E F Shrub layer (3 - 65 - 99) Cormus stolonifera,Lonicera involucrata, VM Rubus spectabilis,Salix lucida,S. sitchensis, Spiraea douglasii W Herb Layer (30 - 60 - 99) AkVA N MA SA Athyrium filix-femina,Equisetum arvense, Lysichiton americanus,Oenanthe sarmen- VW tosa,Scirpus microcarpus Moss Layer (0 - 7 - 50) Soil Moisture Regime pH Mnium spp. St Sl Mo Dy VD Hydrodynamic Index Comments Ws51 swamps often occur between floodplain forests and marshes or shallow-water habitats in flood-scar depressions of larger rivers. Sites are flooded and saturated longer than in the related Ws52

140 Wetlands of British Columbia: A Field Guide to Identiﬁcation Red alder – Skunk cabbage Ws52

Alnus rubra – Lysichiton americanus

General Description Red alder – Skunk cabbage swamps are uncommon in the Georgia Depression and the Coast Mountains at low elevations in small creek draws, ﬂoodplain depressions, and peatland margins. Alnus rubra dominates the canopy. Tall Salix lucida, S. sitchensis, and Thuja plicata are common but of low cover. Rubus spectabilis is usually abundant, and some sites also have abundant Ribes bracteosum or Cornus stolonifera. Lysichiton americanus and Athyrium ﬁlix- femina are prominent in the herb layer; however, some sites have a dominance of Carex obnupta. The moss layer is usually poorly developed because of high litterfall and shading. Humisols or Gleysols with peaty humus forms are most common.

Characteristic Vegetation Wetland Edatopic Grid Tree Layer (0 - .2 - 2) Acer macrophyllum,Alnus rubra, Soil Nutrient Regime Thuja plicata ABCD E F Shrub Layer (15 - 50 - 90) VM Alnus rubra,Cornus stolonifera,Ribes bracteosum,Rubus spectabilis,Salix lucida, S.sitchensis,Sambucus racemosa W Herb Layer (30 - 74 - 99) AkVA N MA SA Athyrium ﬁlix-femina,Carex obnupta, Lysichiton americanus,Maianthemum VW dilatatum,Tiarella trifoliata,Polystichum Soil Moisture Regime pH munitum

Moss Layer (2 - 8 - 35) St Sl Mo Dy VD Eurynchium praelongum,Mnium spp. Hydrodynamic Index

Comments Alnus rubra is a seral species in coastal environments and is also tolerant of lengthy ﬂooding. Cleared Ws53 and Ws54 forests often regenerate to red alder.In these cases,red alder – skunk cabbage forests represent a community successional to conifer forest.Ws52 sites have high watertables and lack elevated microsites that would allow them to succeed to conifer-forested skunk cabbage swamps.

Chapter 5.4 Swamps 141 Ws53 Western redcedar – Sword fern – Skunk cabbage

Thuja plicata – Polystichum munitum – Lysichiton americanus

General Description Western redcedar – Sword fern – Skunk cabbage swamps are uncommon in the Georgia Depression. They occur in receiving sites in topographic depressions, toe slopes, and peatland margins in areas where wet conditions are maintained in hollows, but better-drained sites exist on raised mounds. Thuja plicata dominates the open canopy with Tsuga heterophylla, Acer macrophyllum, and Picea sitchensis. Alnus rubra is also common in natural openings and where clearing has occurred. The shrub layer is moderately developed: Oemleria cerasiformis, Rubus spectabilis, and Sambus racemosa are the most common species. Lysichiton americanus is very prominent in the herb layer along with other rich-site indicators. Gleysol and Humisols are common soil types. Most sites have at least a veneer of dark, woody peat.

Characteristic Vegetation Wetland Edatopic Grid

Tree layer (30 - 70 - 95) Soil Nutrient Regime Acer macrophyllum,Alnus rubra,Picea ABCD E F sitchensis,Thuja plicata,Tsuga heterophylla Shrub layer (10 - 52 - 99) VM Oemleria cerasiformis,Rhamnus purshiana, Rubus spectabilis,Sambucus racemosa W Herb layer (5 - 61 - 95) AkVA N MA SA Athyrium ﬁlix-femina,Dryopteris expansa, Equisetum telmatiea,Lysichiton ameri- VW canus,Maianthemum dilatatum, Polystichum munitum,Stachys chamissonis, Soil Moisture Regime pH Tiarella trifoliata St Sl Mo Dy VD Moss layer (0 - 38 - 80) Hydrodynamic Index Eurhynchium praelongum,Leucolepis acanthoneuron,Mnium spp.

Comments The Ws53 supports moderately productive forest.Soils indicate a wetland environment but plant community composition does not clearly reﬂect this because pronounced microtopography allows upland species to occur.Most species in Ws53 sites are facultative wetland indicators. The Ws53 describes rich,wet skunk cabbage forests of the CDF and very dry CWH; similar site conditions in the rest of the CWH are described by the Ws54. The Ws53 includes site series CDFmm/11 and CWHxm/12.

142 Wetlands of British Columbia: A Field Guide to Identiﬁcation Western redcedar – Western hemlock – Skunk cabbage Ws54

Thuja plicata – Tsuga heterophylla – Lysichiton americanus

General Description Western redcedar – Western hemlock – Skunk cabbage swamps are common in the Coast and Mountains at low elevations. They occur in low-lying areas on ﬂoodplains and receiving sites at toe slopes and wetland margins. These sites are strongly mounded, with conifers on elevated microsites. The canopy is open and consists primarily of Thuja plicata and Tsuga heterophylla. Shrubs root mainly on mounds: Gaultheria shallon, Rubus spectabilis, and Vaccinium species are prominent. Lysichi- ton americanus is always present in damp hollows, accompanied by a diversity of rich- site indicators. Organic veneers of dark, woody humic or mesic peat over ﬁne-textured lacustrine material are typical. However, peat depths are variable, ranging from 0 to 130 cm. Terric Humisols/ Mesisols or Humic Gleysols with peaty humus forms are the most common soil types, but gleyed Podzols also occur. Characteristic Vegetation Wetland Edatopic Grid

Tree layer (0 - 51 - 100) Abies amabilis, Soil Nutrient Regime Picea sitchensis,Thuja plicata,Tsuga ABCD E F heterophylla Shrub layer (4 - 55 - 99) VM Gaultheria shallon,Menziesia ferruginea, Rubus spectabilis,Thuja plicata,Tsuga W heterophylla,Vaccinium alaskaense, AkVA N MA SA V.ovalifolium,V.parvifolium Herb layer (5 - 59 - 99) VW Athyrium filix-femina,Blechnum spicant, Cornus canadensis,Lysichiton americanus, Soil Moisture Regime pH Rubus pedatus,Tiarella trifoliata St Sl Mo Dy VD Moss layer (5 - 63 - 98) Hydrodynamic Index Hylocomium splendens,Mnium spp., Pellia neesiana,Rhytidiadelphus loreus,Sphagnum Group II Comments The Ws54 supports poor to moderately productive forest.Soils indicate a wetland environment but plant community composition does not clearly reflect this because pronounced microtopography allows upland species to occur.Most species in Ws54 sites are facultative wetland indicators.On degrading floodplain sites,forests are dominated by Sitka spruce (see additional units). The Ws54 describes rich,wet skunk cabbage forests of the CWH; similar forests in the CDF and very dry CWH are described by the Ws53.At higher elevations in the MH,forested skunk cabbage ecosystems are described by the Ws55. The Ws54 includes numerous BEC Site Series (see Appendix 4).

Chapter 5.4 Swamps 143 Ws55 Yellow-cedar – Mountain hemlock – Skunk cabbage

Chamaecyparis nootkatensis – Tsuga mertensiana – Lysichiton americanus

General Description Yellow-cedar – Mountain hemlock – Skunk cabbage swamps are common at high elevations in the Coast and Mountains. They occur on toe slopes or depressions with permanent seepage and impeded drainage. The diverse canopy is open and consists of (in descending order of abundance) Chamaecyparis nootkatensis, Tsuga mertensiana, T. heterophylla, and Abies amabilis. The shrub layer is commonly dense with a mix of tree species, Vaccinium spp., Menziesia ferruginea, and Elliottia pyroliﬂorus. Lysichiton americanus is always present in damp hollows, often with abundant Fauria crista-galli. Other herbs common on wet sites frequently occur. The moss layer is a well developed mix of upland and wetland species. Organic veneers of dark, woody humic or mesic peat are typical. However, peat depths are variable, ranging from 10 to 60 cm. Common soil types include Terric Humisols/ Mesisols or Gleysols with peaty humus forms.

Characteristic Vegetation Wetland Edatopic Grid

Tree layer (5 - 38 - 80) Soil Nutrient Regime Abies amabilis,Chamaecyparis nootkaten- ABCD E F sis,Tsuga heterophylla,T. mertensiana Shrub layer (5 - 57 - 95) VM A. amabilis,C. nootkatensis,Elliottia pyro- liﬂorus,Menziesia ferruginea,Rubus W spectabilis,T. heterophylla,Vaccinium AkVA N MA SA alaskaense,V.ovalifolium Herb layer (12 - 62 - 95) VW Blechnum spicant,Coptis asplenifolia, Fauria crista-galli,Lysichiton americanus, Soil Moisture Regime pH Veratrum viride St Sl Mo Dy VD Moss layer (10 - 73 - 95) Hydrodynamic Index Hylocomium splendens,Mnium spp., Rhytidiadelphus loreus,Sphagnum Group II

Comments The Ws55 supports only poor-productivity forest.Soils indicate a wetland environment but plant community composition does not clearly reﬂect this because pronounced microtopography allows upland species to occur.Most species in Ws55 sites are facultative wetland indicators. The Ws55 describes wet skunk cabbage forests of the MH; similar forests at lower elevations in the CWH are described by the Ws54. The Ws55 includes BEC Site Series MHmm1/09,MHmm2/09,MHwh1/09,and MHwh2/09.

144 Wetlands of British Columbia: A Field Guide to Identiﬁcation Additional Swamp Site Associations

Willow – Sedge Salix spp. – Carex spp. Several of the most common Willow – Sedge ecosystems in British Co- lumbia are given full descriptions in this chapter. However, communities dominated by other willow species have been observed in various locali- ties. Some other types observed include those dominated by Salix lucida, S. prolixa, S. glauca, S. planifolia, S. bebbiana, or S. commutata. The reasons for dominance by these other willow species may be the result of several factors: • climatic influences (e.g., S. commutata at high elevations, S. planifolia and S. glauca in cold climates); • special site conditions (e.g., S. lucida and S. prolixa where there is abundant lateral groundwater flow); • willow persistence after hydrological changes (e.g., S. bebbiana); or • stochastic willow establishment. Additional sampling may formalize these communities as distinct Site Associations. Sitka spruce – Skunk cabbage Picea sitchensis – Lysichiton americanus Sitka spruce – Skunk cabbage forests are common in the northern CWH near the edge of the range of western redcedar and uncommon elsewhere along the Coast. They occur on inactive floodplains where drainage is poor and the watertable remains high because of impermeable horizons within the soil profile. However, significant subirrigation and mounded microtophography allow high growth rates of Sitka spruce. The plant community is similar in composition to the Ws54 except that the canopy has little western redcedar and the trees are much larger. Red alder may also occur in the canopy openings. Lysichiton americanus, Rubus spectabilis, Vaccinium spp., Oplopanax horridus, and Athyrium filix-femina are the most common understorey species. In contrast to Ws54 stands, these sites have low cover of Gaultheria shallon, Menziesia ferruginea, and Blechnum spicant. Sitka spruce – Skunk cabbage stands are more productive than the Ws54 swamps that occur more commonly along the Coast. They initiate on active floodplains (high-bench sites) that subsequently become less active and wetter from the formation of restricting layers (duric, placic, ortstein, or clay horizons). Harvesting of Sitka spruce – Skunk cabbage will lead to the site becoming wetter still and sites will likely regenerate as Ws54 in southern areas.

Chapter 5.4 Swamps 145 1

1 Equisetum arvense, common horsetail 2 Fauria crista-galli, deer-cabbage 3 Mnium sp., leafy moss

146 Wetlands of British Columbia: A Field Guide to Identiﬁcation 5.5 SHALLOW-WATERS

1 Water shield mixed with yellow pond-lily in a flooded bog near Parksville, Vancouver Island (CWHxm) 2 A yellow pond-lily community in a lake bay near Meziadin Junction (ICHvc) 3 An alkali pond with Stuckenia pectinata at Lac du Bois grasslands near Kamloops (IDFxh2a)

Chapter 5.5 Shallow-waters 147 SHALLOW-WATERS

Definition Shallow-waters are aquatic wetlands permanently flooded by still or slow- moving water and dominated by rooted submerged and floating-leaved aquatic plants. General Description Vegetation Shallow-waters are dominated by rooted floating-leaved and submerged aquatic plant species and have less than 10% emergent cover. Like marshes, shallow-waters are often simple communities dominated by one to several species. Aquatic environments require adaptations in plants. Anaerobic sediments, light limitations, low carbon dioxide availability, and water currents require specialized structures and metabolic processes. However, in general, these habitats have benign site conditions and are relatively uniform over large climatic regions. Many aquatic plant species in British Columbia occur widely throughout North America. Aquatics typically lack rigid structural components because these are unnecessary for support. Ribbon-like or highly dissected leaves are common in submerged aquatics because they facilitate light penetration and diffusion of dissolved gases. In addition, this leaf arrangement offers little resistance to potentially damaging water movements. Thin leaf cu- ticles facilitate uptake of dissolved gases but this also make species very susceptible to desiccation. Floating-leaved aquatics often have spongy tissue with well-developed air-filled chambers. These “aerhynchema” help transport gases between roots and leaves. Leaves are almost always ovoid and entire to facilitate floating. Pressurized ventilation, a process that moves air from young leaves to roots and back to the atmosphere through older leaves, occurs in many floating-leaved species. Rooted aquatics acquire most of their nutrients from sediments. However, rootless species such as Utricularia spp. and Ceratophyllum spp. absorb nutrients directly from the water. Carbon dioxide diffuses slowly in water and can be limiting for submerged aquatics. Particularly in non-acidic waters, carbon dioxide is converted to bicarbonate. Some species, such as Chara spp., Schoenoplectus subterminalis, Elodea canadensis, and Potamogeton spp., are capable of using

148 Wetlands of British Columbia: A Field Guide to Identification bicarbonate for photosynthesis and often become dominant in stagnant waters where carbon dioxide concentrations are very low. Landscape Position and Distribution Shallow-water ecosystems are always associated with still or slow-moving waterbodies. These sites are widespread and common throughout the province in all climatic regimes. The most common shallow-water habitats occur in the littoral zone of lakes, particularly in protected waters where fine sediments collect, and in potholes. Less commonly, shallow-water ecosystems occur in small peatland ponds, peat degradation hollows, and very sluggish streams. Hydrology and Soils Shallow-water wetlands are affected by factors not well described by the modified edatopic grid. Factors that affect the distribution of aquatic plant species include water chemistry (acidity and salinity gradients), substrate quality, water depth, turbidity, and waterflow. The single most important factor limiting the occurrence of aquatic plants is lack of light. Light levels diminish with increased water depth and turbidity. Turbidity of still waters is largely related to nutrient status of water (Table 5.5.1). The depth to which rooted aquatics occur largely depends on the clarity of the water and can reach as much as 5 m of water where water is very clear. In highly turbid water, light penetration can be limited to several centimetres and only floating-leaved species can occur. Aquatic substrates are generally classified as non-soil because they are permanently flooded at depths greater than 60 cm and do not undergo profile development (Agriculture Canada Expert Committee on Soil

 .. Characteristics of water with different nutrient status (Ellenberg 1986 in Klinka unpublished)

Attribute N and mineral Nutrient Status Water colour Clarity pH availability Dystrophic Yellowish– Very turbid <4.5 Very low deep brown Oligotrophic Greenish– Clear 4.5–7 Low Ca-poor brownish Oligotrophic Blue– Very clear >7 Medium Ca-rich greenish Eutrophic Greenish– Turbid >7 High–Very brownish high

Chapter 5.5 Shallow-waters 149 Survey 1987). Substrates can be sands, silts, clays, muck (a mix of silt, clay, and organic matter), degraded peat sediments, marl, or limnic sediments (gyttja or “loonshit”). Conservation Issues Shallow-water wetlands are among the most important habitats for wildlife and fish. The plants that grow in shallow water are often highly palatable, in part because they lack tough, indigestible structural material. Permanent water and abundant structure encourage use by aquatic macroinvertebrates. Excellent cover and high prey densities attract juvenile and adult fish. Yellow pond-lily is an important forage for Beaver, Muskrat, and Moose. Nutrient loading is potentially one of the biggest impacts on shallow- water ecosystems. High nutrient levels cause blooms of phytoplankton and other algae that reduce water clarity, light penetration, and available oxygen. Many aquatic macrophytes do not tolerate turbid water and will decline. In addition, species that are capable of fast growth under nutrient-loaded conditions, such as Myriophyllum spp., will choke out other species. Sedimentation can affect plant communities by increasing turbidity and decreasing light penetration. Shallow-water Ecosystem Types Shallow-water wetland plant communities have not been well sampled in British Columbia. The descriptions below are based on local classifications, descriptive accounts, and observations. While these summaries provide a scope for discussion and future work, the units presented here should be viewed as preliminary only.

YELLOW POND-LILY TYPES Yellow pond-lily ecosystems are widespread throughout British Colum- bia. They occur in a wide variety of aquatic sites from deep (5 m), clear lakes with gravel substrates to shallow, acidic peat-degradation pools in coastal bogs. Several types have been previously described for British Columbia. Yellow pond-lily - Bladderwort Nuphar lutea – Utricularia macrorhiza This community is widespread on the Coast and in the Interior. It occurs in dystrophic and oligotrophic waters 20–200 cm deep on gyttja and peat

150 Wetlands of British Columbia: A Field Guide to Identification sediments. Sites are relatively species-poor. Nuphar lutea forms an open canopy, with Utricularia macrorhiza and Chara spp. common in the understorey. These communities persist during basin infilling and small patches can be found in bogs. Yellow pond-lily – Water clubrush Nuphar lutea – Schoenoplectus subterminalis This community is widespread in the Georgia Depression in dystrophic and oligotrophic lakes on gyttja and peat sediments. Schoenoplectus subterminalis co-dominates with N. lutea. Utricularia macrorhiza, Najas flexilis, Sparganium natans, and Potamogeton pusillus commonly occur. This community typically occurs in the shallowest shallow-water locations adjacent to bogs or fens. (Described by Ceska 1978.) Yellow pond-lily – Robbin’s pondweed Nuphar lutea – Potamogeton robbinsii This community occurs in the Georgia Depression in 30–120 cm of water on wave-washed shores with gravel bottoms. Vegetation cover is typically low (< 30%). N. lutea forms an open canopy, with Isoetes echinospora, Chara spp., Utricularia macrorhiza, and Najas flexilis in the understorey. (Described by Ceska 1978.) Yellow pond-lily – Richardson’s pondweed Nuphar lutea – Potamogeton richardsonii This community is described by Revel (1972) for the Sub-Boreal Interior. It occurs on mineral sediments with some water movement. N. lutea forms a dense canopy with scattered Potamogeton natans and Polygonum amphibium. Potamogeton richardsonii dominates the understorey; scattered Myriophyllum spicatum also occurs.

PONDWEED TYPES Common pondweed Potamogeton natans Common pondweed communities are widespread in the Interior and the Georgia Depression. They occur in quiet waters on peat sediment in oligotrophic and mesotrophic lakes. They often occur in deeper waters adjacent to yellow pond-lily communities. Potamogeton natans forms a dense canopy. The understorey is frequently sparse; Utricularia spp. and Myriophyllum spp. commonly occur.

Chapter 5.5 Shallow-waters 151 Large-leaved pondweed Potamogeton amplifolius Large-leaved pondweed occurs in the Georgia Depression, Southern In- terior, and Southern Interior Mountains. It occurs in similar conditions to common pondweed but at greater water depths. Light conditions are very limited because of water depth and dense submerged foliage of P. amplifolius; therefore, few other species occur. (Described by Ceska 1978.) Long-stalked pondweed Potamogeton praelongus Long-stalked pondweed occurs throughout the Coast and Interior in deep waters (2.5–4 m) with sandy bottoms. P. praelongus dominates and few other species occur. (Described by Ceska 1978.) Fennel-leaved pondweed – Widgeon-grass Stuckenia pectinata – Ruppia maritima Fennel-leaved pondweed – Widgeon-grass communities occur in saline and alkaline waters of the Central Interior, Southern Interior, and Southern Interior Mountains. Substrates are mineral materials and water depths are 0.5–2.5 m in depth. Ruppia maritima and Stuckenia pectinata both commonly occur in weakly to moderately saline conditions. However, R. maritima is more tolerant of high salinity and will dominate where salinity is above 30 parts per thousand (normal salinity of sea water). Both R. maritima and S. pectinata have very high forage values for waterfowl.

OTHER TYPES Water shield – Bladderwort Brasenia schreberi – Utricularia spp. Water shield ecosystems are common in the Georgia Depression and southwestern Vancouver Island. Conditions are similar to yellow pond- lily sites but water shield usually occurs in deeper waters. Sites are in waters 1–2.5 m in depth with gyttja or peat substrates. Common locations are on the windward site of peatland lakes where sediments and winterbuds accumulate. A dense canopy of Brasenia schreberi is common. Nuphar lutea can occur on some sites, especially in shallower locations. Submerged aquatics include Utricularia gibba, Utricularia macrorhiza, and Ceratophyllum demersum. (Described by Ceska 1978.)

152 Wetlands of British Columbia: A Field Guide to Identification Water smartweed Polygonum amphibium These communties occur in larger lakes in 0.5 –1.5 m deep water on sandy substrates where currents limit accumulation of organic matter and fines. Polygonum amphibium can form a dense floating cover with scattered Potamogeton natans. Submerged species such as Myriophyllum spicatum and Potamogeton foliosus are common. Sites are nitrogen-poor. (Described by Revel 1972.) Water lobelia – Bristle-like quillwort Lobelia dortmanna – Isoetes echinospora Water lobelia – Bristle-like quillwort communities occur in the Georgia Depression and Queen Charlotte Islands in shallow (20–70 cm), oligotrophic waters on sandy- or gyttja-bottomed flats. Vegetation cover is often low (< 30%). Lobelia dortmanna is the dominant species but Isoetes echinospora, Potamogeton gramineus, Ranunculus flammula, and Subularia aquatica are also common. (Described by Ceska 1978.) Muskgrass Chara spp. Muskgrass is a macroalga that occurs in stagnant, alkali waters that have not been over-fertilized or polluted. Chara spp. are efficient at using bicarbonate for photosynthesis and this precipitates large quantities of calcium carbonate (marl). Waterfowl use muskgrass communities extensively. Chara spp. are valuable forage that does not die back in winter and they harbour abundant macroinvertebrates. Wavy water-nymph – Robbin’s pondweed Najas flexilis – Potamogeton robbinsii Wavy water-nymph – Robbin’s pondweed occurs in the Georgia Depres- sion, Southern Interior, and Southern Interior Mountains in clear, fresh to brackish waters, 50–150 cm in depth. Najas flexilis dominates and often has very high cover. Ceska’s (1978) characterization of coastal water-nymph communities includes a high presence of P. robbinsii, P. pusillus, and Utricularia macrorhiza. N. flexilis is an important waterfowl forage species. All parts of the plant are consumed.

Chapter 5.5 Shallow-waters 153 White water-buttercup Ranunculus aquatilis White water-buttercup occurs throughout the province in mesotrophic to eutrophic waters on ﬁrm to soft mineral substrates and often where there is some current. Water depths can be shallow to moderately deep (1.5 m). Narrow-leaved bur-reed Sparganium angustifolium Narrow-leaved bur-reed occurs throughout the province in small ponds and protected embayments. It prefers cold waters 20–100 cm in depth with soft mucky bottoms and non-acid waters. Fruits are eaten by ducks and all parts are grazed by deer and Muskrat.

1 Nuphar lutea, yellow pond-lily 2 Utricularia macrorhiza, greater bladderwort

154 Wetlands of British Columbia: A Field Guide to Identiﬁcation 5.6 ESTUARINE ASSOCIATIONS

1 Khutzeymateen Estuary, northern Coast and Mountains (CWHvm1) 2 Complex of Lyngbye’s sedge and Arctic rush communities on the Bella Coola estuary (CWHms2) 3 Gilttoyees Estuary, near Kitimat, Coast and Mountains (CWHvh2)

Chapter 5.6 Estuarine associations 155 ESTUARINE ASSOCIATIONS

Definition An estuarine ecosystem is an intertidal community, occurring at the confluence of a freshwater source and the marine environment, and is regularly flooded by brackish tidal waters. General Description Vegetation Table 5.6.2 lists species common in Estuarine Site Associations described in this guide. Estuarine Site Associations are characterized by an abundance of species tolerant of repeated (diurnal) flooding and brackish water. Species at the marine edge have specialized morphology to tolerate the desiccating effect of salt water. Succulents and other halophytes with salt-excreting organs are common. Species at the freshwater interface are more typical of wetland habitats but still must be tolerant of at least occasional brackish conditions. In addition, species occurring close to the river course must be tolerant of high sedimentation rates. Low-elevation coastal climates are more equable than interior and high- elevation climates, yet climate still plays a role in estuarine development. Estuaries of the Georgia Depression have “California” species missing from estuaries of the Coast and Mountains ecoregion. Landscape Position and Distribution Estuaries form at the confluence of streams and rivers with the open ocean. The degree of estuarine development depends largely on the size of the river and its sediment load. Larger systems with high sediment loads tend to have extensive estuarine habitat development and diversity. Smaller streams can have substantial estuaries if they are glacier-fed but those systems fed by lakes or peatlands generally have very limited estuarine ecosystem development. Hydrology and Soils Tidal flooding is a characteristic feature of estuarine ecosystems. The lowest vegetated communities (marshes) are flooded and exposed twice daily with each tide. Flood duration is usually several hours per tide cycle. Higher communities are less regularly flooded and flood duration is generally more brief when it does occur (meadows). Soil development is very limited in most estuarine marsh communities. Continual erosion and sedimentation keep soils juvenile. Cumulic

156 Wetlands of British Columbia: A Field Guide to Identiﬁcation Regosols in communities along Salinity the river course are typical. Fresh Oligo- Meso- Poly- Eu- Hyper- Buried vegetation layered between annual spring ﬂood Estuarine Meadows Upper deposits is a common soil feature. Meadows can have some soil development and shallow peat has Estuarine Marshes Wetland – Marshes been observed on some sites. Middle Tidal Zone Tidal Estuarine ecosystems have similar Tidal Flats

characteristics to wetland ecosys- Lower tems with the additional influences of diurnal fluctuations  .. Distribution of ecosystem in watertable and variable salinity classes of estuaries by (Figure 5.6.1). The gradient of elevation and salinity. most importance is the degree of tidal flooding, which is closely related to height above the mean tide level. Ecosystems that occur at the lowest level are flooded with every tide (except neap tides) while the highest may experience only occasional flooding during the highest high tides. The degree of fresh water influence affects species distribution within an estuary independent of other factors. Particularly where high volumes of freshwater flow into estuarine environments, communities change along a gradient from where freshwater influences predominate (usually within the tidal reaches of the river) to where freshwater inputs are minimal. Conservation Issues Estuaries provide critical habitat for many wildlife species but comprise much less than 1% of the coastal landscape. These ecosystems have very high values for waterfowl and shorebirds, which use estuaries to maintain fat stores during migration to the nesting grounds. In addition, estuarine ecosystems are critical for coastal bear populations, providing early-season and mid-season forage, and cover while fishing during the salmon run. Estuaries are critically important fish habitat. Freshwater inputs of nutrients and organic debris into the marine environment fuel highly productive ecosystems. Many saltwater, freshwater, and anadromous fish will selectively use estuaries during some portion of their lifecycle, particularly for juvenile rearing.

Chapter 5.6 Estuarine associations 157 However, river mouths and estuaries are often the only practical access points to resources located farther up the watershed and are a conven- ient location for log booms and camps. Most Estuarine Site Associations are adapted to disturbance and will, therefore, recover from most human-caused physical disturbance except inﬁlling. Damming of the parent river has profound impacts on estuarine ecosystems. Even without changes in water regime, reservoirs settle out ﬂuvial sediments that feed and maintain estuaries. The result is a major reduction in extent of estuarine ecosystems.

Deschampsia cespitosa, tufted hairgrass

158 Wetlands of British Columbia: A Field Guide to Identiﬁcation sq s xx xx xxxx xx xx x xx xxx xxxxx xxx xx x BGPP BWBS SWB ESSF ICH IDF MS SBS CDF CWH SBPS MH Distribution of Estuarine Site Associations by biogeoclimatic zone Widgeon-grass Glasswort – Sea-milkwort saltgrassSeashore Seaside plantain – Dwarf alkaligrass sedge Lyngbye’s sedge – Douglas’Lyngbye’s water-hemlock barley hairgrass – Meadow Tufted hairgrass – Douglas’Tufted aster Alaska plantain rush – Arctic s = southern subzones only q = Queen Charlotte Islands x = incidental; < 5% of estuarine xx = minor; 5–25% of estuarine xxx = major; >25% of estuarine Em01 Em02 Em03 Em04 Em05 Em06 Ed01 Ed02 Ed03  .. 

Chapter 5.6 Estuarine associations 159  .. Estuarine Species Importance Table

Species Em01 Em02 Em03 Em04 Em05 Herbs Ruppia maritima xyzzzz Eleocharis palustris xyz xyz Lilaeopsis occidentalis xyz xy Glaux maritima xyzzz xyzz xyzzz x Salicornia virginica xyzzzz xyzz Distichlis spicata var. spicata xy xyzzzz Spergularia canadensis xy xy xy Atriplex patula xyz x Plantago maritima xyzzz Puccinellia pumila xyzzz Agrostis stolonifera xy Carex lyngbyei xyz xy xyzzzz Potentilla egedii xy xyz Deschampsia cespitosa xxy Triglochin maritima xy xy Juncus arcticus x Plantago macrocarpa Hordeum brachyantherum Angelica lucida Agrostis exarata x Cicuta douglasii Aster subspicatus Conioselinum gmelinii Festuca rubra Lathyrus palustris Ranunculus orthorhynchus Sium suave Lupinus nootkatensis Achillea millefolium x Trifolium wormskioldii Poa trivialis x

160 Wetlands of British Columbia: A Field Guide to Identiﬁcation Em06 Ed01 Ed02 Ed03 Common Name widgeon-grass xyz x x common spike-rush x western lilaeopsis xy x x sea-milkwort American glasswort seashore saltgrass Canadian sandspurry x common orache xy x sea plantain dwarf alkaligrass x creeping bentgrass xyzzzz xyzz xyzz xy Lyngbye’s sedge xyzz xyzz xyzzz xyzzz coast silverweed xyzz xyzzzz xyzzzz xy tufted hairgrass xyz xyz xyz xyz seaside arrow-grass xyz xy x xyzzzz arctic rush xy xy xyzz xyzz Alaska plantain xyz xyz xyzz x meadow barley xyzz x xy xy seacoast angelica xy xy xy xy spike bentgrass xyzz x x Douglas’ water-hemlock xyzz xyzzz xyzzz Douglas’ aster xy x xyz x Paciﬁc hemlock-parsley xy xy xy xy red fescue xyz x xy marsh peavine xyz xy x straight-beaked buttercup xyz x x hemlock water-parsnip xy x x Nootka lupine x xyzz xy yarrow xxyxy springbank clover xy rough bluegrass

Chapter 5.6 Estuarine associations 161 Em01 Widgeon-grass

Ruppia maritima

General Description The Widgeon-grass Site Association is common throughout coastal British Columbia. It occurs in brackish, mud-bottomed pools, lagoons, backwater sloughs, drainage channels, and mudflats that dissect lower portions of estuarine marshes. Tidal inundation is usually prolonged; locations in pools may be permanently flooded. Sites are often small in extent but occasionally can also occur over large areas of tidal flats where sedimentation rates are low. This species-poor community usually consists of pure stands of Ruppia maritima; however, a scattering of other species is possible. Soils are silty Rego-Gleysols. Flooding can be permanent or prolonged during each tidal cycle. Ruppia maritima prefers water depths of approximately 0.5 m but occurs to depths of 4 m.

Characteristic Vegetation Estuarine Grid Tree layer (0 - 0 - 0) Salinity Shrub layer (0 - 0 - 0) Fresh Oligo- Meso- Poly- Eu- Hyper- Herb layer (10 - 30 - 70) Carex lyngbyei,Eleocharis palustris,Lilaeopsis occidentalis,Ruppia maritima Moss layer (0 - 0 - 0) Upper

Comments

Extensive widgeon-grass communities are Middle found on tidal flats with muddy substrates Zone Tidal (protected bays with low wave power or currents).Since Ruppia maritima is tolerant of eusaline conditions,these flats can occur out- Lower side the influence of estuaries.The Em01 can be found in conjunction with any estuarine meadow or marsh ecosystem,in depressions or pools,but it is most commonly below the limit of emergent vascular plants in the middle intertidal. Ruppia maritima also occurs in saline ponds of the Interior (see section 5.8).

162 Wetlands of British Columbia: A Field Guide to Identiﬁcation Glasswort – Sea-milkwort Em02

Salicornia virginiana – Glaux maritima

General Description Glasswort – Sea-milkwort stands are found in the Georgia De- pression and outer coastal areas on sandy or pebbly deposits at the lowest edge of intertidal vegetation. These sites experience daily and prolonged ﬂooding by strongly brackish water. Species diversity is low; typically only Salicor- nia virginica and Glaux maritima are found in abundance. Small patches of Distichlis spicata or Ruppia maritima may occur. Plant cover can be continuous or open. Soils are often ﬁne textured but with a pebbly or gravelly layer that provides better drainage.

Characteristic Vegetation Estuarine Grid Tree layer (0 - 0 - 0) Salinity Shrub layer (0 - 0 - 0) Fresh Oligo- Meso- Poly- Eu- Hyper- Herb layer (15 - 50 - 80) Distichlis spicata,Glaux maritima, Salicornia virginica Moss layer (0 - 0 - 0) Upper

Comments

The Em02 is tolerant of eusaline conditions and Middle may be found outside of estuary inﬂuence in Zone Tidal protected embayments with low wave power.It is often found adjacent to the Em03,which occurs on more poorly drained materials,or the Lower Em01,which tolerates more prolonged ﬂood- ing and continuous soil saturation.

Chapter 5.6 Estuarine associations 163 Em03 Seashore saltgrass

Distichlis spicata var. spicata

General Description Seashore saltgrass sites are found mainly in estuaries of the Georgia Depression but also on southwest Vancouver Island. They occur at the lower limit of estuary vegetation on fine- textured, poorly drained sediments that are flooded for prolonged periods by weakly to strongly brackish water. Species diversity is low. Seashore saltgrass is dominant but there can be significant cover of other low marsh species such as Glaux maritima or Salicornia virginica. Widgeon-grass pools and a scattering of other species sometimes occur. Soils are fine textured and poorly drained.

Characteristic Vegetation Estuarine Grid Tree layer (0 - 0 - 0) Salinity Shrub layer (0 - 0 - 0) Fresh Oligo- Meso- Poly- Eu- Hyper- Herb layer (30 - 70 - 99) Atriplex patula,Distichlis spicata var. spicata, Glaux maritima, Salicornia virginica Upper Moss layer (0 - 0 - 0)

Comments

The Em03 is closely related to the Em02 and Middle these two communities are often found in com- Zone Tidal plex.Hydrology of these two Site Associations is similar but the Em03 occurs on ﬁne-textured material that does not drain when the tide is Lower out.A gravelly or pebbly layer in the Em02 allows for better drainage.

164 Wetlands of British Columbia: A Field Guide to Identiﬁcation Seaside plantain – Dwarf alkaligrass Em04

Plantago maritima – Puccinellia pumila

General Description The Seaside plantain – Dwarf alkaligrass Site Association occurs mainly in the estuaries of the northern Coast and Mountains and protected shores on pebbly or gravelly flats in the middle and upper intertidal. These sites are protected from wave action and often have little freshwater influence. Tidal flooding and exposure occur with most tides. Suitable habitats occur in protected embayments where there is no accumulation of ﬁne-textured sediment; however, such sites appear to be infrequent. Species diversity is low. Em04 sites are often small in extent and consist of a scattered cover of Plantago maritima, Puccinellia pumila, Glaux maritima, and the seaweed, Fucus spp. The substrate is typically sandy/gravelly and gleyed.

Characteristic Vegetation Estuarine Grid Tree layer (0 - 0 - 0) Salinity Shrub layer (0 - 0 - 0) Fresh Oligo- Meso- Poly- Eu- Hyper- Herb layer (30 - 50 - 65) Fucus spp.,Glaux maritima,Plantago maritima,Puccinellia pumila Moss layer (0 - 0 - 0) Upper

Comments

This community is the northern equivalent of Middle the Em02.It occurs below Em05 or even Ed01 Zone Tidal sites where the shore zone is steep.Descriptions of similar communities in Alaska and on Van- couver Island include sites with Puccinellia Lower nutkaensis.No sites with this species have been sampled; however,it is likely that they occur and should be treated as Em04 sites.

Chapter 5.6 Estuarine associations 165 Em05 Lyngbye’s sedge

Carex lyngbyei

The Lyngbye’s sedge Site Association is the most common and widespread estuarine Site Association throughout the coast. It occurs most frequently where there are strong fluctuations of brackish water, active sedimentation, and diurnal flooding and exposure—locations such as tidal flats and channel margins. Low species diversity is typical; Carex lyngbyei often occurs in dense, pure stands. Some sites have scattered Potentilla egedii, Deschampsia cespitosa, Glaux maritima, and Triglochin maritima. Soils are silty or fine-sandy Gleysols or Humic Gleysols. Most commonly they are on sites that experience continual erosion and deposition but do occur on less dynamic sites. Soil profiles frequently exhibit layered mineral deposits with embedded sedge roots and shoots.

Characteristic Vegetation Estuarine Grid Tree layer (0 - 0 - 0) Salinity Shrub layer (0 - 0 - 0) Fresh Oligo- Meso- Poly- Eu- Hyper- Herb layer (30 - 75 - 99) Carex lyngbyei,Eleocharis palustris, Potentilla egedii Moss layer (0 - 0 - 0) Upper

Comments

Along the riverine areas of the estuary,the Middle Em05 is often the lowest vegetated zone.Ed02 Zone Tidal meadows are often directly above Em05 sites in medium-sized estuaries.The related Em06 Site Association replaces the Em05 on sites Lower where freshwater inﬂuences predominate in tidal reaches.

166 Wetlands of British Columbia: A Field Guide to Identiﬁcation Lyngbye’s sedge – Douglas’ water-hemlock Em06

Carex lyngbyei – Cicuta douglasii

General Description The Lyngbye’s sedge – Douglas’ water-hemlock Site Associa- tion occurs in fjord-type estuaries of larger rivers on the northern Coast and Mountains, where large freshwater inputs reduce salinity in the intertidal zone year-round. On the north Coast, the Em06 has been observed along tidal reaches of the Skeena and Nass rivers. It occurs where there are strong fluctuations of weakly brackish water, active sedimentation, and diurnal flooding and exposure—usually in lateral bays along the river. Species diversity is high relative to the similar Em05 that is common in smaller estuaries. Carex lyngbyei is dominant. Many species intolerant of high salinity, especially members of the celery family (Apiaceae) are prominent. Angelica lucida, Cicuta douglasii, Aster subspicatus, and Deschampsia cespitosa are common associates. Soils are always silty or fine-sandy Gleysols or Humic Gleysols that experience continual erosion and deposition. Buried layers of sedge shoots and roots are common in the profile.

Characteristic Vegetation Estuarine Grid Tree layer (0 - 0 - 0) Salinity Shrub layer (0 - 0 - 0) Fresh Oligo- Meso- Poly- Eu- Hyper- Herb layer (80 - 90 - 99) Angelica lucida,Aster subspicatus,Carex lyngbyei,Cicuta douglasii,Deschampsia cespitosa,Eleocharis palustris,Juncus arcticus, Upper Lathyrus palustris,Oenanthe sarmentosa, Potentilla egedii,Ranunculus orthorhynchus, Sium suave,Triglochin maritima

Moss layer (0 - 0 - 0) Middle Tidal Zone Tidal Comments

The Em06 is mostly limited to larger river sys- Lower tems where large freshwater inputs minimize salinity in tidal reaches.Some type locations include the Skeena and Nass rivers. The Em06 can occur adjacent to the Em05, Wm02,or Wm04 sites.

Chapter 5.6 Estuarine associations 167 Ed01 Tufted hairgrass – Meadow barley

Deschampsia cespitosa ssp. beringensis – Hordeum brachyantherum

General Description The Tufted hairgrass – Meadow barley Site Association occurs throughout the Coast in the upper intertidal zone on fan estuaries, on creekside areas within moderate-sized estuaries, and as narrow fringes on steep coastal shores with abundant groundwater seepage. These sites experience daily but generally brief ﬂooding by brackish water. The Ed01 is characterized by relatively low species diversity and a dominance of De- schampsia cespitosa ssp. beringensis. Hordeum brachyantherum occurs commonly and occasionally other grasses, such as Festuca rubra or Agrostis exarata. Potentilla egedii and Carex lyngbyei are often present with low cover. In southern disturbed sites, introduced grasses such as Agrostis stolonifera can be dominant. The soils are usually sandy or loamy-textured Gleysols and Regosols with little or no humus form development.

Characteristic Vegetation Estuarine Grid Tree layer (0 - 0 - 0) Salinity Shrub layer (0 - 0 - 0) Fresh Oligo- Meso- Poly- Eu- Hyper- Herb layer (35 - 65 - 99) Carex lyngbyei, Deschampsia cespitosa ssp. beringensis, Hordeum brachyantherum, Potentilla egedii, Triglochin maritima Upper Moss layer (0 - 0 - 0)

Comments Middle

The Ed01 often occurs above the Em05 and Zone Tidal below forest or Beach dunegrass – Beach lovage ecosystems.This Site Association is similar to the Ed02 but occurs on more saline (and Lower often wetter) sites.

168 Wetlands of British Columbia: A Field Guide to Identiﬁcation Tufted hairgrass – Douglas’ aster Ed02

Deschampsia cespitosa ssp. beringensis – Aster subspicatus

General Description The Tufted hairgrass – Douglas’ aster Site Association is one of the most ﬂoristically diverse and widespread ecosystems in medium to large estuaries in the north and central Coast and Moun- tains. The Ed02 occurs in the high marsh zone between the backshore shrub communities and the low marsh, usually in broad and extensive ﬂats. These sites are limited to zones within the estuary where weakly brackish conditions predominate and inundation is irregular. Deschampsia cespitosa and Aster subspicatus are dominant and diagnostic species, but many other species are often prominent. Soils are mostly Humic Gleysols with silty and sandy textures, but Terric Mesisols have also been encountered.

Characteristic Vegetation Estuarine Grid Tree layer (0 - 0 - 0) Salinity Shrub layer (0 - 0 - 1) Fresh Oligo- Meso- Poly- Eu- Hyper- Herb layer (60 - 85 - 99) Achillea millefolium,Agrostis exarata,Aster subspicatus,Carex lyngbyei,Conioselinum gmelinii,Deschampsia cespitosa ssp.berin- Upper gensis,Hordeum brachyantherum,Plantago macrocarpa, Potentilla egedii,Triglochin maritima

Moss layer (0 - 0 - 0) Middle Tidal Zone Tidal Comments

This community replaces the Em05 as tidal Lower ﬂats accrete and become removed from tidal inﬂuences.

Chapter 5.6 Estuarine associations 169 Ed03 Arctic rush – Alaska plantain

Juncus arcticus – Plantago macrocarpa

General Description The Arctic rush – Alaska plantain Site Association occurs in estuaries of the Georgia Depression and southern Coast and Mountains, where there is a well-developed freshwater lens that reduces the salinity of tidal waters. The Ed03 occurs in the high intertidal zone with brief diurnal tidal inundation—locations that would support Ed01 or Ed02 in more saline environments. Loca- tions can be protected estuaries where fresh water is retained, brackish tidal reaches on larger rivers, estuaries near larger river systems, and localized depressions within estuarine meadows. Juncus arcticus is the site dominant with Plan- tago macrocarpa, Potentilla egedii, and Aster subspicatus common in the herb layer. The herb layer is well developed and displays moderate species diversity of both graminoids and forbs. Silty-textured Gleysols with a humic enriched surface horizon are typical. Minerally enriched Fibrisols also occur.

Characteristic Vegetation Estuarine Grid Tree layer (0 - 0 - 0) Salinity Shrub layer (0 - 0 - 0) Fresh Oligo- Meso- Poly- Eu- Hyper- Herb layer (70 - 80 - 99) Aster subspicatus,Juncus arcticus,Plantago macrocarpa,Potentilla egedii,Triglochin mar-

itima Upper Moss layer (0 - 0 - 0)

Comments Middle

The Ed03 typically occurs above Ed01 or Zone Tidal Em05 sites and at the upper edge of estuarine vegetation before upland ecosystems. Lower

170 Wetlands of British Columbia: A Field Guide to Identiﬁcation Additional Estuarine Site Associations

Canadian sandspurry Spergularia canadensis Canadian sandspurry sites occur in estuaries along the Coast, primarily in the Georgia Depression. They occur on muddy-bottomed depressions in the backshore of estuaries. Sites are protected from waterflow and wave action but have lengthy saturation. A sparse cover of Spergularia canadensis with few other species is typical. Common spike-rush – Lyngbye’s sedge Eleocharis palustris – Carex lyngbyei Eleocharis palustris occurs as a minor component of many estuarine communities throughout the Coast. However, stands dominated by common spike-rush are uncommon and occur in locations with prolonged tidal flooding but low salinity. These can be in protected inlets where there is little tidal flushing or along major river systems where tidal reaches can be fresh water. This Site Association is often represented by a near monoculture of Eleocharis palustris on heavily inundated sites in drainage channels, or with a mixture of meadow species in more protected areas. These latter sites can have Deschampsia cespitosa, Triglochin maritima, Potentilla egedii, and Plantago macrocarpa. The soils are either Gleysols or Organics (in protected sites only). Creeping bentgrass Agrostis stolonifera Agrostis stolonifera is an introduced European species that now forms the dominant component of the high marsh in many estuaries of the Geor- gia Depression. Dune wildrye – Pacific hemlock-parsley Leymus mollis – Conioselinum gmelinii Dune wildrye – Pacific hemlock-parsley communities are common on raised beach ridges or berms where coarse-textured materials have been deposited by beach-forming processes. These sites generally experience little or no flooding. However, salt spray and inundation can occur during storm events. The herb layer is dominated by Leymus mollis. Other species such as Achillea millefolium, Conioselinum gmelinii, Heracleum maximum, and Ligusticum scoticum are often scattered throughout. The tree, shrub, and bryophyte layers are nearly absent. Small flowering plants such as Fritillaria camschatcensis can be found where the site is adjacent to a more protected high marsh community such as Ed02.

Chapter 5.6 Estuarine associations 171 Additional Estuarine Site Associations

Eel-grass Zostera marina Eel-grass beds occur in the upper subtidal areas of estuaries and are exposed only during the lowest low tides. They occur on fine-textured sediments in protected areas where sedimentation rates are not excessive. A open cover of Zostera marina is typical. American bulrush Schoenoplectus americanus American or Olney’s bulrush sites occur in the Fraser River delta and possibly other estuaries of the Georgia Depression in weakly brackish areas below Em05 stands. Schoenoplectus americanus is the site dominant. Soil textures are sandy. Pacific crabapple – False lily-of-the-valley Malus fusca – Maianthemum dilatatum The Pacific crabapple – False lily- of-the-valley Site Association occurs at the upper limit of tidal influence and many sites experience only salt spray and subirrigation. It is described in Chapter 5.7, Flood Classes. Seacoast bulrush Bolboschoenus maritimus Seacoast bulrush sites occur in estuaries of the Georgia Depres- sion in low-lying areas of the high marsh where soils are flooded or at least saturated at low tide and have high salinity. These are often small in extent and a monoculture of Bol- boschoenus maritimus. 1 2

1 Triglochin maritima, seaside arrow-grass 2 Juncus arcticus, arctic rush

172 Wetlands of British Columbia: A Field Guide to Identiﬁcation 5.7 FLOOD ASSOCIATIONS

1 Sandbar willow on wave-washed shore of Nicola Lake (BGxw1) 2 Sitka willow – Red-osier dogwood – Common horsetail low bench on a levee of the Hominka River, east of Prince George (SBSwk1) 3 Low and middle bench communities on the Skeena River floodplain, northern Coast and Mountains (CWHvm1)

Chapter 5.7 Flood associations 173 FLOOD ASSOCIATIONS

Definition A flood ecosystem is a non-wetland ecosystem that occurs on regularly flooded riparian sites with well-drained soils. Sites can be tall shrub (low bench), deciduous forest (middle bench), or coniferous forest (high bench) . General Description Vegetation Table 5.7.2 lists common species of the Low and Middle Bench Site Associations described in this guide. High bench plant community composition is similar to upland seepage sites. High bench associations are not described here because they are thoroughly described in regional  field guides. Low bench ecosystems have a tall shrub structure dominated by willow, alders, and other species tolerant of extended flooding and erosion. Middle benches have similar shrub species but also have a canopy of deciduous trees. The herb layer is dominated by rhizomatous species that can resprout after floods. Even though fresh mineral substrates are frequently exposed, annuals are uncommon on many sites because high shrub cover shades the ground surface. Herb cover is variable because scouring floods and sedimentation temporarily remove above-ground herb growth. These factors also limit bryophyte cover. Landscape Position and Distribution Flood ecosystems occur on the floodplains of rivers or wave-washed lakeshores, where there is deposition of fluvial or lacustrine materials. They usually abut the channel on sites elevated above the mid-season watertable. Lateral bars, midstream bars, point bars, and levees are common site locations for low benches; middle benches often occur on islands, level floodplain benches, and older inner-bend accretion areas. Flood ecosystems occur throughout the province in all zones. However, they are infrequent at higher elevations where there are fewer topographic positions for floodplain development (Table 5.7.1). Hydrology and Soils Flood ecosystems are indundated during the spring freshet in the early part of the growing season. Low benches experience longer (20–40 days) and more powerful flooding than middle benches (< 25 days). Sites can be deeply flooded by stream waters for the first weeks of the growing season but are situated well above normal summer flows. In coastal regions,

174 Wetlands of British Columbia: A Field Guide to Identification fall and winter rain-on-snow Soil Nutrient Regime storms produce the largest annual ABCD E F floods that will affect floodplains M Low and during the dormant season. Middle VM Benches Soils are derived from fluvial sands, gravels, and silts. Vegeta- W AM ANAkVA N MA SA tion grows under aerated soil VW

conditions, with continual subirri- Soil Moisture Regime gation for most of the growing pH season (Figure 5.7.1). Common St Sl Mo Dy VD soil types are Cumulic Regosols of Hydrodynamic Index stratified silts, sands, and gravels.  .. Typically there is no, or very weak, Position of flood ecosystems on the edatopic grid. humus development in low and middle bench ecosystems. Other Comments Flood ecosystems are maintained by a combination of annual flooding, erosion, and deposition. However, most floodplains are geomorphically dynamic; continous sediment deposition, bank erosion, and channel movements modify the site conditions on the floodplain regularly. Middle bench ecosystems will succeed low benches as sites accumulate sediments and become raised above the stream. Continued isolation of middle or low bench ecosystems from the regular flooding (through sediment accumulation or stream channel changes) effectively converts them into seral ecosystems that progress towards high bench or upland ecosystems. Conservation Issues Low and Middle Bench Site Associations occur in the geomorphol- ogically dynamic portion of the floodplain and are maintained by a combination of prolonged flooding and site erosion/sedimentation. The areal extent of flood ecosystems remains constant in a stream reach over time, given no fundamental change in water regime or sediment load, but their location in the floodplain changes in response to stream channel changes. Water control structures reduce the extent of floodplain communities by reducing sediment load and moderating flood levels. Plant species that occur on the active floodplain are tolerant of mech- anical disturbance and will recover from most surface disturbances. Interannual variation in flood intensity and duration is large and most plants will tolerate all but prolonged surface flooding.

Chapter 5.7 Flood associations 175 Flood ecosystems act as sediment traps and prevent rapid erosion of streambank soils by binding soils and slowing floodwaters. In stream systems with high flood power, low bench ecosystems maintain channel form and protect important aquatic habitats. Their riparian location also means that they are major contributors of small and large organic matter that provides nutrients and habitat structure to the stream ecosystem. On small streams, the riparian community provides shade and moderates stream temperature. Flood ecosystems are intensively used by many wildlife species. These are lush habitats with structural elements often not found in adjacent uplands. Adjacency to streams confers additional habitat values to flood sites, including access to water and aquatic food resources (aquatic insects, fish).

Urtica dioica, stinging nettle

176 Wetlands of British Columbia: A Field Guide to Identification xoc x x x xxxx xx xx w w w h BGPP BWBS SWB ESSF ICH IDF MS SBS CDF CWH SBPS MH Distribution of Flood Site Associations by biogeoclimatic zone Mountain alder – Common horsetail alder – Common Mountain fern – Lady dogwood alder – Red-osier Mountain – Horsetail dogwood willow – Red-osier Pacific – Horsetail dogwood willowSitka – Red-osier willow – BluejointDrummond’s Sandbar willow – Rose birch Water – Snowberry – RoseCottonwood dogwood – Red-osier – Spruce Cottonwood club fir – Devil’s – Subalpine Cottonwood x lily-of-the-valley willow – False Sitka alder – SalmonberryRed – Horsetail x alder – Salmonberry – Red Cottonwood xxx x xx x xx xx xx xx x x x xx xx xxx x xx x x x xx x x x xx x xx x xx xx xxx x x x x = incidental; < 5% of flood sites xx = minor; 5–25% of flood sites xxx = major; >25% of flood sites w = wet/very wet subzones only h = warm/hot subzones only xoc = not on outer coast (hypermaritime) Fl01 Fl02 Fl03 Fl04 Fl05 Fl06 Fl07 Fm01 Fm02 Fm03 Fl50 Fl51 Fm50  .. 

Chapter 5.7 Flood associations 177  .. Flood Species Importance Table

Species Fl04 Fl05 Fl06 Fl03 Fl07 Fl01 Fl02 Trees Populus balsamifera ssp. trichocarpa x xyz x xy x Picea X xxxxxxyxy Abies lasiocarpa xx Alnus rubra Picea sitchensis Shrubs Salix sitchensis xyzzzz x x xy Salix drummondiana xyz xyzzzz x Salix exigua x xyzzzz xy Salix lucida xy x xyzzzz x Betula occidentalis xyzzzz Salix bebbiana x x xyz x Alnus incana xy x xy xyzzz xy xyzzzz xyzzzz Cornus stolonifera xyzz x xyzzzz xyz xy xyzz Lonicera involucrata xyz xyzz x xyz xyzz Rosa woodsii xyzz Rosa nutkana xxy Symphoricarpos albus x xyz x Acer glabrum xy Rosa acicularis xxyx xxx Oplopanax horridus xx Rubus parviﬂorus x xxxy Viburnum edule xy x x x Sambucus racemosa x xy xyz Rubus spectabilis Ribes bracteosum xx Herbs Calamagrostis canadensis xy xyzzz x xy xy xy and Equisetum arvense xyzz xy x xyzz xy xyzzz xyzz Dwarf Equisetum hyemale x xyzz x Shrubs Athyrium ﬁlix-femina xy x xyz xyzz Urtica dioica x x xy xyzz Heracleum maximum x xy x xyz xyz Matteuccia struthiopteris x xyzzz Poa pratensis x x xyz x Osmorhiza berteroi xxx Pyrola asarifolia x xxx Actaea rubra xxxy Gymnocarpium dryopteris x xyz xy Circaea alpina xx xxxy Streptopus amplexifolius xxyxy Aster subspicatus x Stachys mexicana Elymus glaucus xxx Maianthemum dilatatum Mosses Brachythecium spp. xy x x xyz xy and Mnium spp. xy xy x x xyz xyz Lichens Rhytidiadelphus squarrosus

178 Wetlands of British Columbia: A Field Guide to Identiﬁcation Fm01 Fm02 Fm03 Fl50 Fl51 Fm50 Common Name

xyzzz xyzzzz xyzzzz x xy xyzzz black cottonwood x xyzz xyzz x spruce x xyz subalpine ﬁr xy xyzzzz xyzz red alder xy Sitka spruce x xyzzzz xy x Sitka willow Drummond’s willow x sandbar willow xx xy Paciﬁc willow x water birch xx Bebb’s willow xy xyzz xyzz x mountain alder xyzz xyzzz xyzz xyzz xyzzz red-osier dogwood x xyz x xy x xyz black twinberry xy prairie rose xyz x x Nootka rose xyzz x x x common snowberry xyz x xy x Douglas maple xy xyz x prickly rose x xyzzzz xyz devil’s club xy xyz xyz xy thimbleberry xyz xyz x xy highbush-cranberry x xyz xy xyz red elderberry x xyz xyzz xyzzz salmonberry xyzz xy stink currant x xy x xyzz x bluejoint xy xyzz xyzz xy xyzzzz xyzz common horsetail xy x x scouring-rush x xyz x xy xy lady fern xxx stinging nettle x xy x xyz xy x cow-parsnip x ostrich fern xyz x Kentucky bluegrass x xy xyz xy mountain sweet-cicely xy xyz xy pink wintergreen x x xyz x x baneberry x xyzz xy xy oak fern x xy xyz xy enchanter’s-nightshade x xyz x clasping twistedstalk x x xyzz Douglas’ aster xyzz x Mexican hedge-nettle x xy xy xyz xy blue wildrye xyzz x xyzz false lily-of-the-valley xxyxx x feathermosses x xyxyxyxy leafy mosses xyz x bent-leaf moss

Chapter 5.7 Flood associations 179 Fl01 Mountain alder – Common horsetail

Alnus incana – Equisetum arvense

General Description Mountain alder – Common horsetail low benches are common throughout the Interior at elevations below 1500 m. They occur on gravel or sand bars adjacent to relatively high-gradient creeks and streams that can have a “flashy” flood regime. Flood events are short during annual spring flooding and occur occasionally during summer storms. Alnus incana is the dominant shrub and forms a continuous canopy on most sites. The understorey can be well developed or sparse depending on recent flood history, but Equisetum arvense usually persists. The moss layer is often very sparse or absent because of high litterfall and recurring sediment deposition. Soils are coarse-textured, often gravelly, Cumulic Regosols and Rego Gleysols.

Characteristic Vegetation Wetland Edatopic Grid Tree layer (0 - 1 - 10) Shrub layer (25 - 75 - 100) Soil Nutrient Regime Alnus incana,Lonicera involucrata ABCD E F Herb layer (1 - 60 - 100) M Athyrium filix-femina,Equisetum arvense, Gymnocarpium dryopteris,Heracleum VM maximum Moss layer (0 - 1 - 40) Brachythecium spp., Mnium spp. W VW Comments Soil Moisture Regime In wetter subzones,Alnus incana stands that occur on fine-textured soils usually have an abundance of Athyrium filix-femina or Mat- teuccia struthiopteris and are described by the Fl02.Alder sites are replaced by willow- dominated Site Associations,such as the Fl05,on lower-gradient streams where fine-textured soils and longer flooding create conditions more favourable to willows.

180 Wetlands of British Columbia: A Field Guide to Identiﬁcation Mountain alder – Red-osier dogwood – Lady fern Fl02

Alnus incana – Cornus stolonifera – Athyrium ﬁlix-femina

General Description Mountain alder – Red-osier dogwood – Lady fern sites are common at low elevations in the wet climates of the Sub- Boreal Interior, Southern Interior Mountains, and Nass Basin, along streams and in creek gullies. Low-gradient floodplains with loamy or fine-textured soils and moderate duration of flooding are characteristic of this Site Association. Alnus incana always dominates the canopy but a diversity of shrubs is common. Cornus stolonifera and Lonicera involucrata are frequently abundant. The understorey is diverse and lush, with a marked abundance of large ferns. In the eastern SBS and in some locations in the Skeena- Nass area, Matteuccia struthiopteris is the dominant fern; elsewhere Athyrium filix-femina predominates. The moss layer is usually very sparse. Soils are Humic Gleysols or Cumulic Regosols.

Characteristic Vegetation Wetland Edatopic Grid Tree layer (0 - 0 - 10) Shrub layer (10 - 78 -100) Soil Nutrient Regime Alnus incana,Cornus stolonifera,Lonicera ABCD E F involucrata,Sambucus racemosa M Herb layer (20 - 75 - 100) Athyrium ﬁlix-femina,Equisetum arvense, VM Heracleum maximum,Matteuccia struthiopteris,Urtica dioica Moss layer (0 - 4 - 20) W Brachythecium spp., Mnium spp. VW Soil Moisture Regime Comments Similar but wetter sites with a perched watertable have an abundance of Lysichiton americanus and are described by the Ws01. Higher-gradient sites with gravelly or sandy soils are occupied by the Fl01.The Fl02 is often found in association with Fm02 or Fm03 middle bench communities.

Chapter 5.7 Flood associations 181 Fl03 Paciﬁc willow – Red-osier dogwood – Horsetail

Salix lucida – Cornus stolonifera – Equisetum

General Description The Pacific willow – Red-osier dogwood – Horsetail Low Bench Site Association is uncommon and widely scattered throughout the Interior and Coast. It has been observed along large, low-gradient rivers with prolonged spring flooding, in locations protected from erosive currents. Salix lucida ssp. lasiandra on these sites can grow to impressive statures and form a closed canopy up to 15 m tall. A sparse to dense cover of Cornus stolonifera, Alnus incana, or Salix prolixa can be present. The understorey is often relatively sparse but horsetails can be abundant. Soils are mostly Regosols derived from deposition of fluvial fine sands and silts. There is little surface organic accumulation.

Characteristic Vegetation Wetland Edatopic Grid Tree layer (25 - 35 - 60) Salix lucida ssp. lasiandra Soil Nutrient Regime Shrub layer (15 - 35 - 40) ABCD E F Alnus incana,Cornus stolonifera, M S. lucida,S.prolixa Herb layer (2 - 17 - 25) VM Equisetum arvense Moss layer (0 - 10 - 40) Mnium spp., Brachythecium spp. W VW Comments Soil Moisture Regime Paciﬁc willow stands are often small in area and dissected by oxbows and drainage channels.They generally occur adjacent to black cottonwood ﬂoodplain ecosystems.More active low bench sites on similar large river systems may be occupied by the Fl06.

182 Wetlands of British Columbia: A Field Guide to Identiﬁcation Sitka willow – Red-osier dogwood – Horsetail Fl04

Salix sitchensis – Cornus stolonifera – Equisetum

General Description Sitka willow – Red-osier dogwood – Horsetail stands are common at low elevations in the wet climates of the Sub-Boreal Interior and Southern Interior Mountains, and in coast transition areas of the Coast and Mountains. They occur primarily on levees or bars in the active floodplains of sluggish, low-gradient streams. Salix sitchensis is consistently the dominant shrub, though some sites have appreciable cover of Alnus incana, Salix drummondiana, or Salix lucida. Equisetum arvense or E. pratense are found on most sites; but where recent floods have deposited new material, the herb layer can be very sparse. Soils are generally fine-sandy, well drained, Cumulic Regosols or Gleysols that remain saturated at depth for much of the growing season.

Characteristic Vegetation Tree layer (0 - 0 - 5) Shrub layer (50 - 95 - 99) Cornus stolonifera,Lonicera involucrata, Salix drummondiana,S. sitchensis Herb layer (1 - 30 - 90) Equisetum arvense Moss layer (0 - 5 - 20)

Comments Wetland Edatopic Grid Wetter sites have sedge- or skunk cabbage– dominated understoreys and are described by Soil Nutrient Regime Swamp Site Associations (Ws06 and Ws51).In ABCD E F drier climates,similar low bench sites are usu- M ally occupied by the Fl05.On the outer Coast, the Fl50 replaces the Fl04.Adjacent middle VM bench communities are Fm50 in coastal areas and Fm02 or Fm03 in interior climates. W

VW Soil Moisture Regime

Chapter 5.7 Flood associations 183 Fl05 Drummond’s willow – Bluejoint

Salix drummondiana – Calamagrostis canadensis

General Description The Drummond’s willow – Bluejoint Low Bench Site Associa- tion is common at lower elevations throughout the Central Interior, Sub-Boreal Interior, and Northern Boreal Mountains, along small, low-gradient streams. Drummond’s willow sites can be deeply flooded during the spring freshet but are much elevated above the mid- season watertable. Salix drummondiana forms a continuous canopy, with other shrubs such as Lonicera involucrata occurring in the understorey. In wetter climates, Spiraea douglasii may co- dominate on some sites. The herb layer has a high cover of Calamagrostis canadensis but is otherwise variably developed, often with open patches of recently deposited fluvial materials. Soils are nearly always silty to fine-sandy textured Cumulic Regosols.

Characteristic Vegetation Wetland Edatopic Grid Tree layer (0 - 0 - 2) Shrub layer (40 - 80 - 99) Soil Nutrient Regime Lonicera involucrata,Salix drummondiana, ABCD E F Spiraea douglasii M Herb layer (4 - 40 - 90) Calamagrostis canadensis VM Moss layer (0 - 1 - 40)

W Comments VW This is the most common Low Bench Site Asso- Soil Moisture Regime ciation on small,low-gradient streams in the sub-boreal forests (SBPS,SBS).It also occurs in the ICH,but in these wetter climates the Fl04 is more common.It also occurs in the BWBS and IDF. Low-lying sites adjacent to the Fl05 are commonly occupied by Ws04 or Wm02.Sites with more powerful ﬂooding,as indicated by coarse sandy and gravelly soils,are often Alnus incana–dominated.

184 Wetlands of British Columbia: A Field Guide to Identiﬁcation Sandbar willow Fl06

Salix exigua

General Description The Sandbar willow Site Association is locally common at low elevations in the Interior along very large river systems. It occurs on sandy lateral bars that receive prolonged spring ﬂooding by powerful currents. In the hot dry subzones of the Southern Interior, sandbar willow sites also occur around large lakes on wave-washed shores. Plant diversity is low. Salix exigua is the site dominant, with a scattering of other species such as Populus balsamifera or Alnus incana appearing with increasing elevation above the stream. Equisetum hyemale is common in the generally very sparse understorey. Especially in warmer climates, annual weeds can seed- in on the exposed mineral soil of these sites. Typically there is no moss layer. Soils are always sandy Cumulic Regosols.

Characteristic Vegetation Tree layer (0 - 0 - 0) Shrub layer (15 - 50 - 80) Populus balsamifera,Salix exigua Herb layer (1 - 5 - 20) Equisetum hyemale Moss layer (0 - 0 - 0)

Comments Wetland Edatopic Grid Salix exigua is a colonial species that resists strong currents,mobile sediments,and pro- Soil Nutrient Regime longed ﬂooding.As sediments accumulate ABCD E F and raise the substrate above the water- M course,Populus balsamifera becomes more competitive and eventually replaces S. exigua. VM Conditions do not appear suitable for S. exigua on smaller river systems.Sufﬁciently W large rivers (such as the Fraser,Thompson, Liard,and Stikine) are uncommon in the Inte- VW rior,limiting the distribution of the Fl06. Soil Moisture Regime

Chapter 5.7 Flood associations 185 Fl07 Water birch – Rose

Betula occidentalis – Rosa

General Description Water birch – Rose ecosystems occur in warm and dry climates of the Southern Interior at low elevations. They occur in the riparian zone of ponds, lakes, and creeks often as a narrow band where flooding is minimal but the watertable remains within the rooting zone for much of the year. Betula occidentalis is consistently a dominant component of the shrub layer, but a variety of other shrub species including Cornus stolonifera, Rosa spp., Salix bebbiana, and Symphoricarpos albus usually occur. The herb layer is often well developed but variable in composition. Soils are fine-textured morainal, lacustrine, or fluvial deposits, often with an organically enriched surface horizon. Gleyed Brunisols and Gleysols are common soil types.

Characteristic Vegetation Wetland Edatopic Grid Tree layer (0 - 10- 30) Betula occidentalis Soil Nutrient Regime Shrub layer (20 - 70 - 85) ABCD E F Betula occidentalis,Cornus stolonifera, M Rosa nutkana,R. woodsii,Salix bebbiana, Symphoricarpos albus VM Herb layer (3 - 35 - 70) Aster spp.,Poa pratensis,Maianthemum stellatum W Moss layer (0 - 1 - 10) VW Soil Moisture Regime Comments Fl07 is different from most Low Bench Site Associations described in this guide; it does occur in classic low bench locations (along watercourses where there is ﬂooding and sedimentation),but also frequently establishes as a fringe habitat around lakes and ponds,where ﬂooding is minimal but the watertable is maintained at depth.

186 Wetlands of British Columbia: A Field Guide to Identiﬁcation Cottonwood – Snowberry – Rose Fm01

Populus balsamifera – Symphoriocarpus albus – Rosa

General Description The Cottonwood – Snowberry – Rose Site Association is uncommon in the dry, warm climates of the Southern Interior and Southern Interior Mountains, where it occurs adjacent to streams, rivers, and lakes on sandy-gravelly ﬂats that are part of the active ﬂoodplain. Flood events are short during the spring freshet and may not occur every year. Populus balsamifera forms an open canopy with a dense to open understorey. A diversity of shrubs is common, with Cornus stolonifera, Symphoricarpos albus, and Rosa species being prominent. The herb layer is variable both in composition and total cover. Most sites have Maianthemum stellatum, Equisetum hyemale, Aster conspicuus, and/or Elymus glauca. Poa pratensis is common on grazed sites. The moss layer is usually absent. Soils are commonly coarse-textured at depth with a loamy or sandy surface horizon. Cumulic Regosols or gleyed Brunisols are typical soil types.

Characteristic Vegetation Wetland Edatopic Grid Tree layer (10 - 35 - 97) Populus balsamifera Soil Nutrient Regime Shrub layer (5 - 45 - 80) ABCD E F Acer glabrum,Amelanchier alnifolia,Cornus M stolonifera,Populus balsamifera,Prunus virginiana,Rosa nutkana,R.woodsii, VM Symphoriocarpus albus Herb layer (1 - 30 - 85) W Elymus glauca,Maianthemum stellatum, Poa pratensis VW Moss layer (0 - 0 - 15) Soil Moisture Regime

Comments These stands reﬂect a different environment than the Fm02,which also occurs in dry climates.The Fm01 has a drier summer water regime because of warm,dry summer climates,shorter,less regular ﬂood period,and coarser,more well-drained soils. Grazing is common in many Fm01 stands in British Columbia and few undisturbed sites occur. The natural herb layer may consist of species such as Maianthemum stellatum,Equisetum hyemale,Aster conspicuus,and Elymus glauca,with Poa pratense increasing on grazed sites. The Fm01 includes several existing BEC Site Series (see Appendix 4).

Chapter 5.7 Flood associations 187 Fm02 Cottonwood – Spruce – Red-osier dogwood

Populus balsamifera – Picea X – Cornus stolonifera

General Description The Cottonwood – Spruce – Red-osier dogwood Site Associa- tion is the most common middle bench community of low elevations thoughout the Interior on suitable sites. It occurs on sandy or gravelly fluvial materials adjacent to streams and rivers with short flood durations followed by continual subirrigation. Populus balsamifera forms an open canopy with scattered interior spruce. Cornus stolonifera and Alnus incana are dominant in the shrub layer, but frequently with some cover of Viburnum edule, Rosa acicularis, and Lonicera involucrata. Along smaller river systems, C. stolonifera is often sparse and A. incana dominates. The herb layer can be well- developed or sparse depending on recent flood history, but Equisetum arvense usually persists. The moss layer is always poorly developed. Soils are Cumulic Regosols or Gleyed Brunisols.

Characteristic Vegetation Wetland Edatopic Grid Tree layer (10 - 40 - 80) Picea X,Populus balsamifera Soil Nutrient Regime Shrub layer (12 - 60 -100) ABCD E F Alnus incana,Cornus stolonifera,Lonicera M involucrata,Picea X,Populus balsamifera, Rosa acicularis,Viburnum edule VM Herb layer (1 - 30 - 75) Equisetum arvense Moss layer (0 - 2 - 50) W VW Contents Soil Moisture Regime The Fm02 describes middle bench communities from a wide range of climatic zones.The overwhelming influence of flood effects limits the species composition to those that can tolerate flooding. The Fm01 includes several existing BEC Site Series (see Appendix 4).

188 Wetlands of British Columbia: A Field Guide to Identiﬁcation Cottonwood – Subalpine ﬁr – Devil’s club Fm03

Populus balsamifera – Abies lasiocarpa – Oplopanax horridus

General Description The Cottonwood – Subalpine fir – Devil’s club Site Associa- tion is uncommon in the cold interior rainforest climates of the Nass Basin and Sub-Boreal Interior. It occurs on sandy or gravelly flats adjacent to streams and rivers with relatively prolonged flood durations. Annual spring flood events are short during the freshet but there is prolonged subirrigation. Cottonwood forms an open canopy with scattered subalpine fir and interior spruce. Oplopanax horridus is consistently abundant in the understorey. Cornus stolonifera, Sambucus racemosa, and Alnus incana frequently occur. The herb layer is often moderately developed with abundant Gymnocarpium dryopteris, Equisetum arvense, Athyrium filix-femina, and other forbs. Soils are sandy or gravelly Cumulic Regosols or Gleysols.

Characteristic Vegetation Wetland Edatopic Grid Tree layer (15 - 30 - 80) Abies lasiocarpa,Picea X,Populus Soil Nutrient Regime balsamifera ABCD E F Shrub layer (48 - 80 - 99) M Abies lasiocarpa,Alnus viridis, A. incana, Cornus stolonifera,Oplopanax horridus, VM Picea X,Sambucus racemosa,Viburnum edule Herb layer (5 - 45 - 95) Actaea rubra,Athyrium ﬁlix-femina, W Dryopteris expansa,Equisetum arvense, VW Gymnocarpium dryopteris,Osmorhiza Soil Moisture Regime berteroi,Pyrola asarifolia,Rubus parviﬂorus, Streptopus amplexifolius Moss layer (0 - 1 - 60) Rhytidiadelphus loreus

Comments The related Fm02 occurs on sites with more prolonged soil saturation and in regions with warmer summer climates.

Chapter 5.7 Flood associations 189 Fl50 Sitka willow – False lily-of-the-valley

Salix sitchensis – Maianthemum dilatatum

General Description The Sitka willow – False lily-of-the-valley Site Association is uncommon in the Coast and Mountains, where it is restricted to floodplains of maritime climates. It is generally found at the transition between the freshwater conditions of the fluvial system and the uppermost reaches of brackish influence in estuaries. Sitka willow sites can experience brief or temporary annual floods during the spring freshet but are much elevated above the mid-season watertable. The shrub layer is dominated by Salix sitchensis, often with little development of other shrub species. The herb layer is moderately well developed and supports Maianthemum dilatatum and Calamagrostis canadensis. Other graminoid species and forbs such as Agrostis exarata, Aster subspicatus, and Sanguisorba canadensis are common. The moss layer is often poorly developed. Soils are nearly always loamy to sandy-textured Gleysols or Regosols.

Characteristic Vegetation Wetland Edatopic Grid Tree layer (0 - 0 - 3) Shrub layer (45 - 70 - 90) Soil Nutrient Regime Rubus spectabilis,Salix sitchensis ABCD E F Herb layer (15 - 47 - 65) M Agrostis exarata,Angelica genuﬂexa,Aster subspicatus,Calamagrostis canadensis, VM Heracleum maximum,Maianthemum dilatatum,Sanguisorba canadensis Moss layer (5 - 15 - 35) W Rhytidiadelphus squarrosus VW Soil Moisture Regime Contents In outer coastal areas,the Fl50 replaces the Fl04 of more inland climates.Fl51 can be found in similar but slightly drier and better- drained sites.Tidal effects on soil moisture regime probably favour willows,which are more tolerant of prolonged ﬂooding of the rooting zone.

190 Wetlands of British Columbia: A Field Guide to Identiﬁcation Red alder – Salmonberry – Horsetail Fl51

Alnus rubra – Rubus spectabilis – Equisetum arvense

General Description Red alder – Salmonberry – Horsetail low benches are widespread in the Coast and Mountains. They occur adjacent to river courses where ﬂood duration is lengthy and sedimentation is abundant. Alnus rubra forms a closed tall shrub or low tree canopy. Cornus stolonifera, Ribes bracteostum, and Rubus spectabilis are prominent in the understorey. The herb layer can be sparse or well-developed depending on recent ﬂood history. Equisetum arvense always persists but other species commonly occur. The moss layer is often very sparse. Soils are typically sandy Cumulic Regosols.

Characteristic Vegetation Tree layer (0 - 9 - 50) Alnus rubra Shrub layer (15 - 70 - 95) Alnus rubra,Cornus stolonifera,Ribes bracteosum, Rubus parviﬂorus,R. spectabilis,Sambucus racemosa Herb layer (2 - 17 - 50) Circaea alpina,Elymus glauca,Equisetum arvense, Stachys mexicana Moss layer (0 - 0 - 10)

Comments Wetland Edatopic Grid Cleared high and middle bench floodplain forests will often regenerate to A. rubra and Soil Nutrient Regime in these cases will represent a community ABCD E F successional to conifer forest.Fl51 stands es- M tablish on sites with more lengthy flooding than Fm50 but also on similar sites where VM stand-initiating floods occur in autumn (a common occurrence in coastal watersheds). A. rubra drops seed in fall and will establish W quickly on exposed mineral soils.Populus bal- VW samifera drops seed in spring and its seedling Soil Moisture Regime will not establish where a thick cover of red alder already exists. The Fl51 includes several existing BEC Site Series (see Appendix 4).

Chapter 5.7 Flood associations 191 Fm50 Cottonwood – Red alder – Salmonberry

Populus balsamifera – Alnus rubra – Rubus spectabilis

General Description The Cottonwood – Red alder – Salmonberry Site Association is common along rivers in the Coast and Mountains. River benches that are ﬂooded annually for moderately long periods are typical. Populus balsamifera dominates the canopy but a subcanopy of Alnus rubra and scattered conifers is typical. The shrub layer is well-developed, often with Rubus spectabilis and Cornus stolonifera both being prominent. The herb layer can be sparse or well-developed, depending on recent ﬂood history and cover of the canopy. Equisetum spp. and Maianthemum dilitatam are the major constituents. The moss layer is generally poorly developed. Soils are sandy Cumulic Regosols.

Characteristic Vegetation Wetland Edatopic Grid Tree layer (20 - 67 - 90) Alnus rubra,Picea sitchensis,Populus Soil Nutrient Regime balsamifera,Thuja plicata ABCD E F Shrub layer (15 - 70 - 95) M Alnus rubra,Cornus stolonifera,Lonicera involucrata,Oplopanax horridus,Picea VM sitchensis,Rubus spectabilis,Sambucus racemosa Herb layer (3 - 45- 95) W Equisetum arvense,Maianthemum VW dilatatum Soil Moisture Regime Moss layer (0 - 1 - 30)

Comments The typical floodplain successional sequence in the inner Coast and Mountains is Fl06 >> Fm50 >> Sitka spruce – Salmonberry forests.These communities often occur adjacent to each other,indicating the accumulation of fluvial sediments,which progressively raises benches higher above normal waterflow. Fm50 sites with silty or clayey layers within the soil profile are common.These sites often have some species more indicative of wetter sites such as Oenanthe sarmentosa,Carex utriculata, or Scirpus microcarpus. The Fm50 includes several existing BEC Site Series (see Appendix 4).

192 Wetlands of British Columbia: A Field Guide to Identiﬁcation Additional Flood Site Associations

Green alder Alnus viridis Green alder is a common component of well-drained but periodically disturbed sites, such as avalanche tracks. Riparian fringe communities of Alnus viridis occur on river systems where flood periods are very short and soils are well-drained or where ice scour occurs during spring break- up. These are mostly narrow bands adjacent to streams. The understorey is variable. Pacific crabapple – False lily-of-the-valley Malus fusca – Maianthemum dilatatum The Pacific crabapple – False lily-of-the-valley Site Association occurs on the outer Coast at the upper limit of tidal influence in the transition between the upland forest and estuarine ecosystems. Inundation occurs, generally briefly and often during the spring freshet, when salinity is low. However, many sites experience salt spray and tidal subirrigation. Malus fusca is the site dominant, accompanied by a sparse to well developed and diverse forb-dominated understorey. Picea sitchensis can be present on raised microsites with limited tidal influence. On floodplains, Pacific crabapple can progress to Sitka spruce forests because sediments accumulate and the site is raised higher above floodwaters. Paper birch – Red-osier dogwood Betula papyrifera – Cornus stolonifera Fluvial stands dominated by paper birch (Betula papyrifera) are occasionally encountered where the Fm02 is expected. These stands have shrub and understorey species are generally similar to the Fm02. The primary reason for the dominance of paper birch on these sites is likely the occurence of a stand-initiating fall flood. Unlike cottonwood, which seeds in the early summer as spring floods are receding, paper birch seeds in the fall. A fall flood that exposes mineral soil and then does not flood the following spring will regenerate to B. papyrifera. Most paper birch–dominated sites are in cool subzones or landscape positions (e.g., at the base of north-aspect slopes), which also seems to favour paper birch over cottonwood. Red-osier dogwood Cornus stolonifera In southern areas of the province, low bench communities characterized by dense thickets of red-osier dogwood occur adjacent to small streams. On some of these sites, other shrub species such as Alnus incana or

Chapter 5.7 Flood associations 193 Additional Flood Site Associations

Betula occidentialis also occur. Dense shrub layers often limit herbaceous growth. Prolonged flooding occurs on many sites. Soils are typically fine- textured and poorly drained. Trembling aspen – Red-osier dogwood Populus tremuloides – Cornus stolonifera Trembling aspen – Red-osier dogwood forests occur in drier climatic areas throughout the Interior where sites have seepage or subirrigation but limited flooding. They occur in riparian locations as well as upland habitats. Suitable riparian areas include lake edges and terraces of streams. Trembling aspen forms an open to closed canopy. Cornus stolonifera, Symphoricarpos albus, and Rosa acicularis form a well- developed and sometimes very dense shrub layer. Herb and moss layer composition and development is variable but Equisetum arvense is often prominent.

1 Cornus stolonifera, red-osier dogwood 2 Calamagrostis canadensis, bluejoint 3 Equisetum arvense, common horsetail

194 Wetlands of British Columbia: A Field Guide to Identiﬁcation 5.8 “TRANSITION” ASSOCIATIONS

1 Saline meadow zonation at the Meadow Lake marshes, 100 Mile House (IDFdk3) 2 A lush high-elevation Barclay’s willow – Arrow-leaved groundsel shrub-carr, Causqua Creek (ESSFwv) 3 Dried lake bed with abundant Salicornia rubra, Stinky Slough near Cranbrook (IDFdm2)

Chapter 5.8 “Transition” associations 195 “TRANSITION” ASSOCIATIONS

Two classes of ecosystems, saline meadows and shrub-carrs, have traditionally been described as wetlands in British Co- lumbia (Runka and Lewis 1981; Steen and Roberts 1988). Neither of these ecosystem types meets the soils or vegetation critieria for wetlands. However, these ecosystems frequently occur adjacent to wetlands in a zone transitional to upland ecosystems and have structural similarities. For this reason, several of the more common saline meadow and shrub-carr site associations are presented here. The environmental factors that cause the occurrence of these two classes are different but, for purposes of this guide, they are informally treated together as a “Transition” group. The saline meadow class is a member of the Grassland Group of the Terrestrial Realm; the shrub-carr class is a member of the Shrubland Group of the Terrestrial Realm. Saline meadows Saline meadows are ecosystems with moist, saline or alkaline soils that a) occur within the drawdown zone of shallow temporary or permanent ponds and lakes and b) are dominated by salt-, alkali-, and inundation-tolerant graminoids and forbs. Vegetation Table 5.8.2 lists species common to “transition” site associations. Saline meadows have a distinctive ﬂora dominated by halophytes. Most sites are grass-, rush-, or sedge-dominated but extremely saline environments support only succulents. In British Columbia, there are no trees or shrubs species tolerant of saline soils. A group of diminutive moss species occurs almost exclusively in saline meadows (McIntosh 1986). In many cases, saline meadows are dominated by congeners of estuarine species. These include the following saline meadow vs. estuarine pairs: Deschampsia cespitosa ssp. cespitosa vs. D. cespitosa ssp. beringensis, Dis- tichlis spicata var. stricta vs D. spicata var. spicata, Hordeum jubatum vs. H. brachyantherum, Juncus balticus vs. J. arcticus, and Potentilla anserina vs. P. egedii. Landscape Position and Distribution Saline meadows occur primarily in the drawdown zone of small waterbodies, where there is early-season saturation or shallow ﬂooding that give way to generally well-aerated soil conditions for much of the growing season. They are common in warm, semi-arid climates of the South

196 Wetlands of British Columbia: A Field Guide to Identification and Central Interior but mostly Soil Nutrient Regime absent elsewhere in the province ABCD E F (Table 5.8.1). M Shrub Saline Carrs Meadows Hydrology and Soils VM Sites can be shallowly inundated or merely saturated in the early W AM ANAkVA N MA SA season; thereafter, the watertable VW falls well below the surface Soil Moisture Regime (Figure 5.8.1). Evaporation of pH standing water accumulates salts. St Sl Mo Dy VD Soils are usually loamy or fine- Hydrodynamic Index textured Gleyed Brunisols,  .. Position of “transition” classes Humic Gleysols, or Solenetzics. on the edatopic grid. Humus is thin. Shrub-carrs Shrub-carrs are low shrub ecosystems that occur on moist mineral soils in areas prone to growing-season frosts, which would support forested ecosystems under normal circumstances. Vegetation Shrub-carrs are always low shrub cover types and usually have highly diverse herb and moss layers. Few obligate hydrophytes occur. Shrubs that are tolerant of growing-season frosts and few growing degree days dominate. Landscape Position and Distribution Shrub-carrs occur in frost-prone depressions or cold-air drainage valleys where frost and cold, moist soils preclude establishment of trees. Such sites are often at the edge of wetlands but shrub-carrs can also occupy entire basins. Shrub-carrs are most common in the cold and dry climates of the Chilcotin Plateau, the western Fraser Plateau, and the Northern Boreal Mountains. Hydrology and Soils Sites have at most very moist soils and are never flooded. They are fed by groundwater or lateral subirrigation from adjacent wetlands. Soils are imperfectly drained and cold. Distinctly hummocky microtopography is common.

Chapter 5.8 “Transition” associations 197 Soils are typically Gleyed Brunisols or Gleysols with thin moder or mull humus forms. Conservation Issues Both saline meadows and shrub-carrs occur in ranching country. These sites often have good forage and grazing potential. Forage production coupled with riparian location means that these sites often receive heavy use by livestock. Species such as Deschampsia cespitosa and Puccinellia nuttalliana will be replaced by Poa pratensis and Hordeum jubatum, respectively, under heavy grazing pressure. Saline meadows have a limited distribution in the province and occur in areas where wetlands are even more important than usual for wildlife. The red-listed Tiger Salamander commonly rears in alkali waterbodies of the extreme southern Interior and will use the adjacent saline meadow as adults.

1 2

1 Carex praegracilis, field sedge 2 Distichlis spicata var. stricta, alkali saltgrass 3 Hordeum jubatum, foxtail barley

198 Wetlands of British Columbia: A Field Guide to Identiﬁcation v v v v x xx xx xx d dc dc d x xx xx xx d d d d dc dc x x dc BGPP BWBS SWB ESSF ICH IDF MS SBS CDF CWH SBPS MH Distribution of “Transition” Site Associations by biogeoclimatic zone Alkali saltgrass barley alkaligrass – Foxtail Nuttall’s Field sedge hairgrass Tufted – KinnikinnickScrub birch moss – Glow willow Grey-leaved groundselArrow-leaved willow – Barclay’s x xx xxx xx xx x xx xx x xx xx x = incidental; < 5% of wetlands xx = minor; 5–25% of wetlands xxx = major; >25% of wetlands d = dry subzones only v = dry subzones of the SBPS only dc = dry and cold subzones only Gs01 Gs02 Gs03 Gs04 Sc01 Sc02 Sc03  .. 

Chapter 5.8 “Transition” associations 199  .. “Transition” Species Importance Table

Species Gs01 Gs02 Gs03 Gs04 Shrubs Salix brachycarpa Betula nana Salix glauca x Salix barclayi Herbs Distichlis spicata var. stricta xyzzzz xy x Spartina gracilis xyzz xy Suaeda calceoliformis xyz x Aster ericoides ssp. pansus xyz xy x Poa secunda xy x x Hordeum jubatum xyzz xyzzz xy Puccinellia nuttalliana xyz xyzzzz Carex praegracilis xy xyz xyzzzz Elymus trachycaulus xxxyx Poa pratensis x x xyz xy Aster ericoides xxy Potentilla anserina xxxy Juncus balticus xy xy xyzz xy Deschampsia cespitosa xy xyzzzz Potentilla gracilis xy x Taraxacum ofﬁcinale xxyxy Carex utriculata xyz Achillea millefolium xxy Muhlenbergia richardsonis x Kobresia myosuroides Koeleria macrantha x Arctostaphylos uva-ursi x Antennaria pulcherrima Maianthemum stellatum xx Aster ciliolatus x Calamagrostis canadensis xxyxy Thalictrum occidentale xx Fragaria virginiana xx Senecio triangularis Valeriana sitchensis Epilobium angustifolium Erigeron peregrinus x Sanguisorba canadensis Trollius albiﬂorus Equistem arvense Mosses Bryum pseudotriquetrum xy Drepanocladus spp. xy xy Aulacomnium palustre x Brachythecium spp. xxy Mnium spp. xx

200 Wetlands of British Columbia: A Field Guide to Identification Sc01 Sc02 Sc03 Common Name xyzz x short-fruited willow xyzzzz xyzzz scrub birch xyz xyzzzz grey-leaved willow x x xyzzzz Barclay’s willow alkali saltgrass alkali cordgrass seablite x tufted white prairie aster x Sandberg’s bluegrass x foxtail barley Nuttall’s alkaligrass xyz x x field sedge xy x slender wheatgrass xy x Kentucky bluegrass x tufted white prairie aster x common silverweed xyzz xy Baltic rush xxy tufted hairgrass xy xy graceful cinquefoil xy x common dandelion xy x beaked sedge xyz xy yarrow xyzzz mat muhly xy Bellard’s kobresia xy junegrass xyzzz xyz kinnikinnick xyz x showy pussytoes xy x star-flowered false Solomon’s-seal xy xyz Lindley’s aster x xyz xyz bluejoint xy xyz x western meadowrue xyz xyzz x wild strawberry x xyzz arrow-leaved groundsel xyzz Sitka valerian x xyz xy fireweed xxysubalpine daisy xy Sitka burnet xy x globeflower xyz common horsetail x xyz xy xxy hook-mosses x xyzzz xyzz glow moss x xyz feather-moss xyz leafy mosses

Chapter 5.8 “Transition” associations 201 Gs01 Alkali saltgrass

Distichlis spicata var. stricta

General Description The Alkali saltgrass Saline Meadow Site Association is uncommon in the BG, PP, and dry IDF of the Central Interior and Southern Interior at elevations below 1000 m. Gs01 meadows occur in the seasonally flooded riparian zone of small potholes and shallow lakes where evaporation accumulates salts. Brief flooding in the early season is followed by pronounced surface drying, occasionally leaving a distinct salt crust. Only salt-tolerant plants are found on these sites; no shrubs or mosses occur. Distichlis spicata var. stricta is always prominent but some sites have high cover of Spartina gracilis, Amphiscirpus nevadensis, or Poa secunda ssp. juncifolia. Soils are fine textured, saline or saline-alkali, imperfectly drained materials with minimal organic accumulation. Solonetzes and Gleysols are common soil groups.

Characteristic Vegetation Wetland Edatopic Grid Tree layer (0 - 0 - 0) Shrub layer (0 - 0 - 0) Soil Nutrient Regime Herb layer (22 - 82 - 92) ABCD E F Amphiscirpus nevadensis,Aster ericoides ssp. M pansus,Distichlis spicata var. stricta, Hordeum jubatum,Puccinellia nuttalliana, VM Salicornia rubra,Spartina gracilis,Suaeda calceoliformis W Moss layer (0 - 0 - 0) VW Comments Soil Moisture Regime Sites occur that have a high abundance of Spartina gracilis,Amphiscirpus nevadensis, or Poa secunda ssp.juncifolia.These sites are currently considered variations of the Gs01; further sampling might support separation of these ecosystems into new Site Associations. Gs01often occurs adjacent to shallow open-water sites and in complex with Gs02 sites.Sites that are highly saline are often dominated by Sueda calceoliformis or Salicornia rubra. Hordeum jubatum is a naturally occurring species on Gs01 sites but becomes more prominent with grazing or mineral soil exposure. This Site Association was previously described as part of a Saltgrass – Alkaligrass Wet Meadow Site Association by Steen and Roberts (1988).

202 Wetlands of British Columbia: A Field Guide to Identiﬁcation Nuttall’s alkaligrass – Foxtail barley Gs02

Puccinellia nuttalliana – Hordeum jubatum

General Description Nuttall’s alkaligrass – Foxtail barley saline meadows are uncommon in the dry IDF, MS, and SBPS subzones of the Central Interior and Southern Interior at elevations between 800 and 1200 m. Gs02 meadows occur in the seasonally flooded riparian zone of small alkali potholes and shallow lakes where evaporation accumulates salts. Brief flooding in the spring gives way to merely moist conditions during the summer. High overall graminoid cover is common; Puccinellia nuttalliana is a constant dominant. Hordeum jubatum occurs naturally with low cover on most sites but increases and may become dominant with soil disturbance. Shrubs are absent and the moss layer is poorly developed. Soils are often fine textured, alkali, or saline-alkali Gleysols or Solonetzs on poorly to imperfectly drained materials. Sites often have dark surface horizons.

Characteristic Vegetation Wetland Edatopic Grid Tree layer (0 - 0 - 0) Shrub layer (0 - 0 - 0) Soil Nutrient Regime Herb layer (40 - 85 - 90) ABCD E F Carex praegracilis,Hordeum jubatum, Puccinellia nuttalliana M Moss layer (0 - 0 - 10) VM

Comments W Gs02 can occur alone in basins or adjacent to VW other saline meadows or marshes such as the Soil Moisture Regime Wm07 or Wm06.It occurs at generally higher elevations than the Gs01.In the IDF of the Chilcotin Plateau,where both Gs01 and Gs02 are relatively common,the Gs02 occurs on more alkali sites. The Gs02 was previously described as part of a Saltgrass – Alkaligrass Wet Meadow Site As- sociation by Steen and Roberts (1988).

Chapter 5.8 “Transition” associations 203 Gs03 Field sedge

Carex praegracilis

General Description Field sedge meadows are common throughout the Chilcotin Plateau region of the Central Interior and uncommon in the Southern Interior at elevations below 1250 m. Gs03 sites form extensive stands in seasonally ﬂooded, moderately alkaline depressions; or peripheral communities in the drawdown zone around permanent ponds and Wm06 or Wm07 marshes. The Gs03 occurs where there is brief early-season inundation followed by a dropping of the watertable below the surface. The upper horizons often dry out by the early growing season. Carex praegracilis is the constant dominant on these sites. Juncus balticus occurs on wetter examples but never dominates (see Wm07). On grazed sites Poa pratensis becomes prominent but will occur even on undisturbed sites. These sites usually have no shrub and little moss layer development. Soils are ﬁne textured slightly alkaline Gleysols or gleyed Brunisols developed in poorly to imperfectly drained lacustrine materials, with up to 10 cm of surface organic accumulation.

Characteristic Vegetation Wetland Edatopic Grid Tree layer (0 - 0 - 0) Shrub layer (0 - 0 - 0) Soil Nutrient Regime Herb layer (60 - 90 - 99) ABCD E F Carex praegracilis,Juncus balticus, M Poa pratensis Moss layer (0 - 10 - 20) VM

Comments W Gs04 commonly occurs adjacent to the closely VW related Wm07,which occupies wetter loca- Soil Moisture Regime tions.Interannual variation in water depth is typical where these Site Associations occur, and the extent of Gs03 may increase during drier years.The Gs03 occurs on less saline sites than the Gs02. The Gs03 was previously described as part of a Baltic rush – Field sedge Wet Meadow Site Asso- ciation by Steen and Roberts (1988).

204 Wetlands of British Columbia: A Field Guide to Identiﬁcation Tufted hairgrass Gs04

Deschampsia cespitosa ssp.cespitosa

General Description Tufted hairgrass meadows are uncommon in the cold, dry subzones of the Central Interior (SBPS and MS). They form extensive communities in frost-prone basins fed by seepage from the surrounding upland. These sites are usually saturated to the surface in the early part of the growing season. Deschampsia cespitosa ssp. cespitosa can form nearly pure stands and gives the site a tussocky appearance. Shrub and moss layers are poorly developed. In wetter microsites Carex utriculata can be prominent. Soils are often ﬁne textured, imperfectly drained, and weakly alkaline Brunisols or humic Gleysols with up to 15 cm of surface organic accumulation.

Characteristic Vegetation Wetland Edatopic Grid Tree layer (0 - 0 - 0) Shrub layer (0 - 0 - 0) Soil Nutrient Regime Herb layer (25 - 85 - 99) ABCD E F Carex utriculata,Deschampsia cespitosa M Moss layer (0 - 20 - 90) VM Comments Gs04 can occupy entire shallow depressions W but more commonly it occurs in the moist ri- VW parian area around Wm01 marshes or small Soil Moisture Regime ponds.In some areas,it may also be in complex with dry meadows dominated by Danthonia intermedia or Festuca altaica. Deschampsia cespitosa is widely distributed in the province and is a common dominant in alpine and coastal estuarine ecosystems. The Gs04 is described by Steen and Roberts (1988).

Chapter 5.8 “Transition” associations 205 Sc01 Scrub birch – Kinnikinnick

Betula nana – Arctostaphylos uva-ursi

General Description The Scrub birch – Kinnikinnick Shrub-carr Site Association is common in the colder, drier subzones of the Central Interior. These shrub-carrs form small communities in frost-prone basins with moist, cold substrates and often surround larger wetlands. In drier climates, these sites are rarely, if ever, inundated, but subsurface saturation is typical in the early season. Sites are distinctly mounded with shrubs on relatively dry organic-rich mounds. The Sc01 has very high species diversity. Betu- la nana dominates the shrub layer with high cover of Salix brachycarpa and S. glauca. Arctostaphylos uva-ursi and Muhlenbergia richardsonis are common dominants of the very diverse herb layer. The moss layer is poorly developed and variable. Soils are often ﬁne textured, poorly to imperfectly drained materials with thin surface organic accumulation. Gleysols and gleyed Brunisols are common soil types.

Characteristic Vegetation Wetland Edatopic Grid Tree layer (0 - 0 - 0) Shrub layer (10 - 58 - 80) Soil Nutrient Regime Betula nana,Salix brachycarpa,S.glauca ABCD E F Herb layer (40 - 80 - 99) M Achillea millefolium,Antennaria pulcherrima, Arctostaphylos uva-ursi,Carex praegracilis, VM Fragaria virginiana,Juncus balticus,Muhlen- bergia richardsonis W Moss layer (0 - 15 - 40) VW Comments Soil Moisture Regime Sc01 occurs alone in shallow depressions or around the periphery of Wf01,Wm01,or Gs03 ecosystems.Though the Sc01 and Sc02 occupy similar frost-prone sites,the Sc01 occurs on drier site conditions. Betula nana–dominated ecosystems are widespread in the Boreal,especially at higher elevations in the SWB.However,few plots in these communities have been established; it is possible that the Sc01 also occurs in the Northern Boreal Mountains.Other scrub birch–dominated Shrub-carr Site Associations certainly occur but remain undescribed. The Sc01 is described by Steen and Roberts (1988).

206 Wetlands of British Columbia: A Field Guide to Identiﬁcation Grey-leaved willow – Glow moss Sc02

Salix glauca – Aulacomnium palustre

General Description Grey-leaved willow – Glow moss shrub-carrs are uncommon in the colder, drier subzones of the Interior from the Southern Interior to the Northern Boreal Mountains. They form small communities in frost-prone basins and hollows with moist, cold substrates fed by seepage from upslope sites. These sites are often wetter than the Sc01. Standing water is not present between mounds, and subsurface saturation may be common early in the growing season. Salix glauca grows on elevated mounds and dominates the shrub layer. The herb layer is diverse with large numbers of species all having sparse cover. The moss layer is often well developed. Soils are often ﬁne textured, poorly to imperfectly drained materials with up to 15 cm of surface organic accumulation.

Characteristic Vegetation Wetland Edatopic Grid Tree layer (0 - 0 - 1) Shrub layer (20 - 80 - 90) Soil Nutrient Regime Betula nana,Salix glauca ABCD E F Herb layer (10 - 50 - 90) M Arctostaphylos uva-ursi,Aster ciliolatus,Cala- magrostis stricta,C.utriculata,Deschampsia VM cespitosa,Epilobium angustifolium,Fragaria virginiana,Thalictrum occidentale,Valeri- ana dioca W Moss layer (10 - 50 - 99) VW Aulacomnium palustre,Bryum pseudotri- Soil Moisture Regime quetrum

Comments Though the Sc02 and Sc01 occupy similar frost-prone sites,the Sc02 occurs on moister site conditions and at higher elevations.Salix glauca–dominated ecosystems are widespread in the Boreal,especially at higher elevations in the SWB.However,few plots in these communities have been established; it is likely that additional Grey-leaved willow Site Associations occur. Sc02 is locally common at montane to low subalpine elevations (1070–1615 m) in the BWBS, MS,SBPS,and SWB zones. The Sc02 is described by Roberts (1984).

Chapter 5.8 “Transition” associations 207 Sc03 Barclay’s willow – Arrow-leaved groundsel

Salix barclayi – Senecio triangularis

General Description The Barclay’s willow – Arrow-leaved groundsel Shrub- carr/Swamp Site Association is common in the subalpine climates of the Interior from the Southern Interior to the Northern Boreal Mountains. These shrub-carrs form extensive communities on subalpine seepage slopes, gullies, abandoned stream flats, and lake margins with cold, moist to very moist (wet) soils. Standing water is typically not present, but subirrigation is common. Salix barclayi is always present but can be shorter in stature than surrounding forb species. The herb layer is diverse, well developed, and dominated by subalpine forbs such as Senecio triangularis and Valeriana sitchensis. Soils are commonly fine to medium textured, poorly to imperfectly drained mineral materials with well-humified surface organic horizons. Gleysols and gleyed Brunisols are most common but occasionally these sites occur on shallow peat.

Characteristic Vegetation Wetland Edatopic Grid Tree layer (0 - 0 - 0) Shrub layer (10 - 80 - 90) Soil Nutrient Regime Salix barclayi,S.commutata ABCD E F Herb layer (5 - 50 - 90) M Calamagrostis canadensis,Equisetum arvense,Senecio triangularis,Valeriana VM sitchensis Moss layer (10 - 55 - 99) W Aulacomnium palustre,Brachythecium spp., Mnium spp. VW Soil Moisture Regime Comments Sc03 is very widespread in British Columbia and can occupy substantial area in plateaus of upper montane and subalpine forest lands.It occurs alone in extensive ﬂats or associated with forb meadows and high-elevation fens.The Wf04 has a similar structure and occurs over the same geographic range but develops under wetter conditions.

208 Wetlands of British Columbia: A Field Guide to Identiﬁcation Conservation and 6 management issues

Wildlife uses of wetlands are as diverse as wetland types.While marshes and shallow waters support the highest wildlife populations, bogs and other wetland types each support their own distinctive wildlife communities. Wetlands and related ecosystems provide ecosystem services dispropor- tionate to their limited extent in the landscape. They are valuable ecosystems with many functions: • the majority of wildlife and ﬁsh species in the province use these ecosystems for part of their life history and some are entirely dependent upon them; • they support large populations of economically important fur-bearers; • wetlands and related ecosystems provide some of the highest-quality range for livestock; and • they are an integral component of hydrological systems and good water quality. This chapter outlines some general management principles for wetlands and related ecosystems. In addition, important habitat attributes and life history requirements for wetland-using wildlife are presented.

LIVESTOCK MANAGEMENT

Wetlands (mainly fens and marshes) and related ecosystems are important grazing lands throughout much of British Columbia. They are productive ecosystems often with a diversity and abundance of palatable species that maintain their forage quality later in the season than adjacent uplands. In addition, livestock favour riparian habitats because of their association with drinking water, cool microclimates, and shade. However, these characteristics can lead to habitat overuse and degradation by livestock without proper management. Some effects of improper livestock management systems in riparian areas include: • damage, reduction, or elimination of vegetation by browsing, grazing, or trampling; • changes in plant communities through selective browsing and grazing; • soil compaction and disturbance, which increases erosion and de- creases water availability to plants; • changes in ﬂuvial process and aquatic ecosystems, through bank shearing, reduction of vegetative cover, and subsequent changes to stream channel characteristics; • decrease in water quality though increased water temperatures, nutrients, suspended sediments, and bacterial counts; and • reduced wildlife habitat quality from impacts to vegetation structure, species composition, and water quality.

210 Wetlands of British Columbia: A Field Guide to Identification A full review of livestock effects on riparian areas in British Columbia is presented in Powell et al. (2000). Range Conditions The Range Resources Assessment Manual (1999) outlines procedures for identifying proper functioning condition () of riparian areas. One attribute for assessment of range condition is the “potential natural community” () (B.C. Ministry of Forests 2002). The Site Association descriptions in Chapter 5 can be used as guidelines for identifying the . A management plan that targets the maintanence of a healthy  will often limit impacts to other important indicators of  such as: • Stream channel shape and bank stability: Excessive livestock use of streams and riparian areas can lead to widening and shallowing, stream trenching, or braiding, depending on the texture of fluvial materials and level of use. • Vegetation structure: Some ecosystems, particularly willow swamps and low-bench ecosystems, can undergo structural changes (such as removal of low shrub layers, resprouting, clubbing, and highlining) that are indications of reduced ecosystem functioning. • Soil structure: Sites with wet and fine-textured soils are susceptible to compaction and rutting, which will lead to an increase in erosion and weedy species. • Water nutrient levels: Large inputs of livestock feces can elevate nutrient levels in streams and wetlands, leading to eutrophication. • Wildlife: Wildlife can suffer direct impacts through trampling of ground nests and indirect impacts through habitat degradation and displacement. Strategies for Minimizing Livestock Impacts on Vegetation Livestock impacts on vegetation can be minimized through proper con- trols on timing and level of use. There are, broadly, four methods of managing livestock in riparian areas. 1) Control animal distribution: This generally means fencing to limit access to sensitive areas. However, providing dedicated access points or alternative sources of water, forage, and shelter away from wetlands and streams can reduce overuse of riparian zones. 2) Control timing of access: Negative impacts of livestock on native plant species are often related to season of use. Early-season use coincides with the period of highest resource demands by growing plants. Livestock use should be timed to occur after spring when key plants are less sensitive. For example, willows are adapted to and

Chapter 6 Conservation and management issues 211 tolerant of winter browsing but appear to be susceptible to early growing-season browsing. The advantages of late-season grazing include good plant vigour and productivity, minimal soil disturbance, and minimized disturbance to wildlife during the breeding season (Kauffman 1982). Many wetland sites have fine-textured soils that are susceptible to degradation when wet. Early-season use of wet sites by livestock can result in soil compaction, rutting, and erosion. 3) Provide adequate rest periods: Season-long, continuous use will have detrimental effects on range condition. Range programs with long growing-season rest periods allow plants to acquire and store resources, produce seed, and establish seedlings. Most native plants are not adapted to continual grazing and will be replaced by non- native species that are either unpalatable for livestock or tolerant of heavy use. Rest-rotation systems that allow for 1 rest year out of 3 or 2 rest years of spring-summer grazing out of 3 have been recommended to pre- serve riparian functioning (Kauffman and Krueger 1984). 4) Control grazing intensity: It is estimated that utilization rates below 25% will show little effect on streamside vegetation, but rates over 65% typically show measurable impacts (Platts 1982). Degraded sites may not have to be rested to allow full recovery if grazing intensity is low (Manoukian and Marlow 2002). However, where wildlife browsing is heavy, even light browsing by livestock can reduce productivity of willow (Brookshire et al. 2002).

Cattle will heavily use riparian areas around wetlands but use marshes mainly to access drinking water.

212 Wetlands of British Columbia: A Field Guide to Identiﬁcation TIMBER HARVEST

Of the Site Associations described in this guide, forest harvesting is possible only in some forested swamps (Ws07, 08, 10, 11, 53, 54, 55), bogs (Wb08), and middle-bench stands; other ecosystems do not have trees large enough for commercial timber production. Specific management guidelines can be found for most of these ecosystems in the regional  field guides. All wetlands are potentially affected by adjacent land clearing, especially where such activities are widespread or involve potential disruption of groundwater flow. Harvesting Flood Forests Functioning riparian areas are very important for stream ecosystems. Streamside vegetation provides the following important functions: • maintains bank stability and channel form by binding soils and trap- ping sediments; • provides large woody debris to streams, which produces important fish habitat; • inputs fine organic debris that fuels stream ecosystems; • moderates surface water temperatures; • provides habitat for terrestrial and aquatic wildlife; and • maintains in-channel and off-channel fish habitat. Issues in Floodplain Management Of the flood ecosystems in this guide, only cottonwood-dominated middle- bench communities are harvested commercially.2 Most commercial sites occur on the floodplains of large rivers where extensive stands can be ex- ploited for pulpwood. See Petersen et al. (1996) Black Cottonwood and Balsam Poplar Managers’ Handbook for British Columbia for issues and approaches to management of these stands. Red alder stands can also be commercial but it is primarily successional forests that are managed. Bank and floodplain erosion: Removal of tree cover can lead to instability of the floodplain and stream banks during flood events. Channel avulsion will be more common where vegetation has been removed. Silvicultural issues: Prolonged flooding and vigorous competing brush can cause regeneration difficulties. Loss of seedlings can be an important issue in years when rodent populations are high.

2 High-bench ﬂoodplain forests, where commercial timber harvesting is common, are not covered in this guide.

Chapter 6 Conservation and management issues 213 Wildlife trees: Large cottonwood are valuable wildlife trees while standing or fallen. Large old cottonwood will be selected by birds that build large platform-like nests (e.g., Osprey, Bald Eagle, Great Blue Heron) and primary and secondary cavity nesters. They are valuable for several listed species such as Western Screech Owl, Lewis’ Woodpecker, and various bat species. Some level of mature canopy retention is recommended on cottonwood floodplains outside of the reserve zone. Fish/Forestry: Trees within one tree height of the water are particularly important for stream ecosystems. They provide shading, bank stability, litterfall, and large woody debris. Along coastal streams, floodplains provide critical off-channel habitat for coho salmon (Oncorhynchus kisutch). Temporary ponds in riparian forest are used extensively by over-wintering coho juveniles. This habitat needs to be assessed during winter high flows when it is most readily identified. Several management practices can minimize impacts (Hartman and Brown 1988): • place culverts and bridges to permit movement of fish; • maintain natural drainage; • fall and yard away from wet depressions; and • do not treat the riparian zone as a sediment trap; keep sedimentation to a minimum. Harvesting Swamp and Bog Forests Draining marginally productive swamp forest to improve tree growth is a common practice in the boreal forest outside of British Columbia and may play an increasing role here as timber reserves diminish. Most forested swamps are between intermediate wetland and upland habitats and will present significant silvicultural challenges if harvested. • Sensitive soils. Wet and fine-textured soils are highly susceptible to compaction and rutting. • Overly wet site conditions. Good tree growth is often limited to raised microsites. Where harvesting removes much of the canopy, watertables often rise, further reducing microsites that are dry enough to support trees. • High capability for competing vegetation. Successful reforestation will require brush control. • Frost-prone locations. Many wetlands occur in low areas of the landscape where cold air ponds and growing-season frosts are common. Frost damage and seedling mortality can be expected in cleared areas with no overstorey cover. • Low site potential. The site index at 50 years for forested wetlands is less than 12 m for bogs and less than 17 m for most swamps. However,

214 Wetlands of British Columbia: A Field Guide to Identification this likely overestimates site potential for many of these sites for two reasons: 1) Severe site limitations restrict the maximum size of trees on these sites; height growth curves begin to flatten as early as 50 years. 2) Sampling methods for site index focus on the best growing-site trees; tree growth on wetland sites, however, is highly microsite- dependent with the largest trees growing on a few optimal microsites. Total volume production on forested wetlands is thus very low. Characteristics of Sites Where Timber Harvest Should Be Avoided There are several indicators that can be used to identify forested swamps that should probably not be harvested. 1) Sites with poor tree growth. These sites have low timber values and will clearly be the most problematic for reforestation. Site preparation such as mounding or ditching will likely be required to create drier microsites for planting. However, these site preparation tech- niques can result in further site degradation by creating extensive water ponding that restricts tree root development. 2) Some sites with good tree growth occur on deep organic soils with abundant lateral seepage. While the mature timber values may be good, they will prove difficult to reforest. All of the problem issues listed above will need to be considered. Only winter harvesting, while wet soils are frozen, will be possible. Mounded microtopography is typically critical to good tree growth on these sites. Heavy machinery will often damage this structure even in winter. Many elevated microsites are not composed of mineral soil but created from old “tip-ups” where replanting success is unlikely. Mounding site treatments are generally not useful on these sites due to high erosion rates of well-humified organic materials. Advanced regeneration can be critical for successful re-establishment of a coniferous canopy (Päivänen 1997). Cluster planting on raised microsites may be neces- sary to ensure adequate survival and growth of conifers. 3) Sites with standing water in summer. Trees in swamps extract abundant groundwater for growth, removing vast quantities of soil water and lowering the watertable through evapotranspiration. The presence of standing water on the site during the growing season indicates that the site is near the lowest threshold of productive tree growth. Removal of the canopy will lead to deeper and more persistent surface water; the ability of trees to regenerate will be compromised and availability of plantable spots will be further limited.

Chapter 6 Conservation and management issues 215 Harvesting around Wetlands Impacts of harvesting upland forests in the riparian zone on adjacent wetland plant communities is typically not marked unless road building activities interrupt water inflow or outflow, or if forest clearing is extensive and affects overall watershed hydrology. Where the riparian zone is composed of swamp forest at level with the wetland, removal of large portions of it will lead to a raising of the watertable (see above), which can have an impact on plant communities in the rest of the wetland. Most wetland-using wildlife species use both the wetland and adjacent riparian area. When viewed from a wildlife perspective, the wetland and its riparian area should be viewed as a single functioning wetland ecosystem. Terrestrial habitats may be exceptionally important for conservation of some species groups such as amphibians. Management plans targeting amphibian conservation that focus only on wetland land habitats are likely to be unsuccessful without consideration of the riparian area (Marsh and Trenham 2001). Harvesting that maintains some canopy cover, minimizes soil disturbance and compaction, and retains downed wood will minimize impacts to wildlife populations. Several objectives should be considered when harvesting around wetlands: • Protect wetland hydrology – Ensure that surface and groundwater inlets and outlets are not interupted by road construction or harvesting activities. • Maintain vegetative cover – Implement partial-cutting harvest plans and windfirm edges. • Maintain wildlife trees – Maintain large standing snags and veterans. • Maintain downed wood – Leave large stems on the ground. • Avoid wet depressions – Avoid harvesting or skidding through low- lying areas of the stand. • Winter harvest – Harvest during season of lowest wildlife use. Landscape Planning The cumulative effects of resource extraction, urban development, and transportation networks have resulted in a disconnected landscape. Noss (1994) cites Shaffer’s (1992) assertions that long-term viability of large carnivores in North America is being compromised by the fragmenta- tion of the original wilderness into small refugia. This phenomenon may be occurring at smaller scales for less mobile wetland animals, as wet-

216 Wetlands of British Columbia: A Field Guide to Identiﬁcation lands become isolated from each other by development activities and resource extraction. Semlitsch and Bodie (1998) argue that small wetlands are extremely valuable for maintaining diversity in a number of plant, invertebrate, and vertebrate taxa (e.g., amphibians). However, the ability of many amphibian species to disperse may be under-appreciated and may be limited only where signiﬁcant barriers, such as roads or urban development, exist (Marsh and Trenham 2001).

WETLANDS AS WILDLIFE HABITAT

Wetlands and related ecosystems have variable wildlife values. Some basic features that may influence a wetland’s wildlife habitat value include: 1. Presence of water. Aside from the obvious habitat values for aquatic invertebrates, fish, and amphibians, ponds, lakes, and streams are often focal points for terrestrial animals as primary sources of drinking water. 2. Structural diversity and cover. In many landscapes, wetlands and riparian ecosystems are structurally distinct from the surrounding upland and provide unique habitats. High structural complexity within wetlands generally increases their value to wildlife by providing nesting cover and foraging habitat for a wide range of species. 3. Abundant forage. Wetlands and riparian areas can be productive habitats. Many wetland plant species provide important forage. Skunk cabbage, sedges, and horsetails form an important component of bear diets; pond-lilies, willows, and other shrubs are browsed by moose; and marsh emergents and aquatics are foraged by Muskrat, Beaver, and waterfowl. 4. High prey densities. The productive aquatic habitats adjacent to flood communities and shallow-water habitats frequently produce high concentrations of aquatic insects that are the food base for larger animals, especially birds and bats. 5. Unique habitat. Peatlands provide unique microhabitats that are used by specialized invertebrates (Finnamore and Marshall, 1994). 6. Rarity in the landscape. Especially in drier climates, water and aquatic habitats are especially valuable for wildlife but are also uncommon and often in areas of development. Good-quality wetland habitat is at a premium in these areas. The most heavily used wetlands are marshes, with their high productivity and adjacency to open water, followed by swamps, fens, and bogs.

Chapter 6 Conservation and management issues 217 However, all of these habitats are vital for wetland-dependent species and important for upland species that use wetlands and their associated riparian areas for food, water, and cover. Amphibian Habitat Requirements Amphibians are generally associated with wetland habitats or other moist environments (excluding saline habitats, which are toxic to amphibians). British Columbia has 20 amphibian species, many of which are restricted to the relatively mild climates of the Coast and southern Interior. Adults of some amphibian species stay in or near water always; others stay near water but feed in adjacent uplands; several of British Columbia’s amphibians are mainly terrestrial, using water only for breeding; and some are wholly terrestrial (Table 6.1). In British Columbia, all amphibians, except Plethodontid salamanders, require water in which to lay their eggs. Wetlands are the most common habitats but some species such as Clouded Salamander and Tailed Frog lay their eggs in streams. Others such as Spadefoot Toad will deposit eggs in temporary ponds. Broadly, wetlands with good vegetation structure for the egg mass attachment and sufficient flood duration for juvenile development are preferred breeding sites. Frogs generally metamorphose during the same season that the eggs are laid while salamanders can take several years to mature, especially in cold climates. In a study of amphibian distribution in the Puget Sound Basin, Richter and Azous (1995) found that there was no statistically significant relationship between amphibian richness and wetland size, number of vegetation classes, presence of predators, characteristics of waterflow, and wetland permanence. Low-velocity flow and low water-level fluctuation were correlated with high species richness. Increasing water-level fluctuation and percent watershed urbanization were correlated with low species richness. Amphibians are especially vulnerable to environmental pollutants. Their breathable skins readily absorb chemicals. Clearly, additions of pesti- cides, fuel, or other chemicals to waterbodies will affect local populations of amphibians and should be avoided. Precipitous declines in some amphibian populations have been observed around the world. The cause of these declines has still not been pin- pointed but several causal factors are likely, including increased ultraviolet (-) irradiation, acid precipitation, adverse weather pat-

218 Wetlands of British Columbia: A Field Guide to Identification prefers shallow water with water vegetation shallow prefers ponds and and lakes and low-elevation High-  grassland ponds; lakes in cold neotenes Feb–AprilFeb–AprilMar–May in burrows along shores also common not necessarily permanent streams bodies and slow-moving Red List;Red Okanagan on CoastCommon April–May water Migrates to Common; widespread breed to water Migrates to S. of 56°N. as on Coast breed Common to water Migrates to in burrows Underground Blue List; Okanagan breed ponds to Migrates to forests Moist (e.g., microhabitats Moist Locally common, breed to water Migrates to widespread but patchy early as December Small, alkali, often and temporary lakes on CoastCommon elevation Sept–Oct Forests Migrates May–June; at highLocally common; for breeding water Migrates to NE B.C. During dry weather, under the soil Locally common;SW B.C. ponds (alkali) or shallow Temporary for breeding water Migrates to habitats in all biogeo- forested Most Forests; and shrubs; on trees often List;Red and pastures rubble) in forests rock southern and streams; Lakes also in subalpine Interior or permanent pools and Temporary aquatic; Mainly and ecoregions zones climatic other to move may Common, widespread in ponds; in pools along streams Meadows, with water lots of Shallow deciduous forests, and vegetation; Interior small ponds in damp forests Near aquatic; ponds or swamps Migrates to Very land over move may Mar–JuneScattered; in common northern B.C. edges of At ponds and lakes small ponds; small ponds Mar–April breeds or permanent ponds and standing water Shallow Temporary Van. Introduced for breeding water to Is. Moves and fringes of sites, breeding Near Vegetated also may but permanent water- SW mainland Mar–June aquatic; Mainly May–July breeds Van. Introduced Is. and SW mainland aquatic; Mainly Shallow, April–July periods during permanent rainy marshes, May–July breeds near ponds; and forest Meadows ponds, edge permanent ponds and small lakes and Prefers Water edge Water clear ponds for only a few days Shallow marshes around and fields forage in meadows alpine tundra; tolerant very cold ponds of Permanent variable depth; lakes, especially with emergent vegetation ponds Permanent streams; slow-moving Mar–April in north in early spring Distribution and habitat use by adult pond-breeding amphibians (from Province of British Columbia 1998) Species Distribution Seasonal habitat Terrestrial habitat Aquatic Tiger Salamander Northwestern Salamander Long-toed Salamander Rough-skinned Newt Basin Great Spadefoot Toad Western Treefrog Pacific Striped Chorus Frog Red-legged Frog Northern Leopard Frog Frog Spotted Frog Wood American Bullfrog Frog Green  . 

Chapter 6 Conservation and management issues 219 terns, environmental pollution, or infectious disease. While most of these are beyond the scale of management by field workers, the latter should be carefully considered by individuals who are visiting several wetlands. In North America, mortalities caused by amphibian iridoviruses or Ranaviruses have been documented for species that occur in British Columbia, including Leopard Frog, Red-legged Frog, and Tiger Sala- mander. These viruses can persist under adverse conditions (such as dried mud on boots) for several months to several years. Field workers should, at minimum, rinse outerwear and equipment. Transport of adults or juveniles from one locality to another should also be avoided. Pond-breeding Amphibians The Rough-skinned Newt is the only newt species in British Columbia. It occurs throughout the Coast. Adults forage for slugs and worms in open seral and mixed forests near permanent water. Adults also feed on tadpoles and aquatic invertebrates. Shallow water in swamps, fens, and bogs is used for breeding. Larvae are carnivorous. Four species of mole salamander occur in British Columbia: Long-toed Salamander, Northwestern Salamander, Tiger Salamander, and Pacific Giant Salamander. Adult mole salamanders spend much of their adult life underground in rodent burrows or under rocks. Juveniles are always aquatic and some are neotenous. Long-toed Salamander are widespread on the Coast and in the Interior mostly south of 56 degrees north latitude. Adults prefer forested edge habitats near water and breed in still waters with abundant aquatic vegetation. Northwestern Salamander occur along the Coast and are frequently neotenous. Larvae, if they develop into terrestrial adults, often take more than a single year to metamorphose. Breeding occurs in fishless, permanent, shallow ponds. Tiger Salamander are widespread in North America but occur only in the warmest, driest areas of the southern Interior (south Okanagan) in British Columbia. Adults live mainly underground in grassland habitats. Breeding is primarily in alkali ponds and shallow lakes. Larvae transform in 3–4 months and live primarily in warm areas of ponds with abundant algae.

220 Wetlands of British Columbia: A Field Guide to Identification Great Basin Spadefoot occur in the dry climates of the Southern Interior. This is primarily a terrestrial species but eggs are laid in temporary or shallow alkali lakes and ponds. Larvae eat detritus, carrion, and aquatic vegetation and mature in about 6 weeks, after which they leave the natal pond. The Western Toad occurs throughout British Columbia in all but the coldest climates. This widespread species is primarily terrestrial, travel- ling far from open water and wetlands. They prefer moist habitats where they can escape desiccating conditions. Breeding occurs in shallow ponds and pools with sandy bottoms. Two species of treefrog occur in British Columbia: Pacific Treefrog and Northern Chorus Frog. The former is widespread along the Coast, Southern Interior, and Southern Interior Mountains. The latter occur only in the Boreal Plains and Taiga Plains. Pacific Treefrog adults forage in forest edges and shrubby habitats sometimes far from open water. Shrub swamps and shrub fens are good habitats. Breeding occurs in shallow, weedy, permanent shallow water. Northern Chorus Frog live mainly underground in grassy or wooded areas. Breeding occurs anywhere there is shallow water. Four native frog species occur in British Columbia: Red-legged, Spotted, Northern Leopard, and Wood Frog. These “true” frogs are the most aquatic of the frog groups in British Columbia and are much more closely associated with open water as adults. Red-legged Frog occur on Vancouver Island and the adjacent mainland where they occur in small waterbodies in forested landscapes. Adults can wander some distance into forest but most commonly remain near water. Spotted Frog are widespread throughout the Interior in cold-water ponds and lakes. Northern Leopard Frog are found in the Southern Interior Mountains and Southern Interior around permanent waterbodies. Adults forage in grassy uplands often far from open water. Eggs are laid in small, well- vegetated ponds and swamps.

Chapter 6 Conservation and management issues 221 Wood Frogs commonly use peatlands adjacent to open water.

Woodfrog occur throughout the Interior, mostly north of 51 degrees north latitude. This species is very cold-tolerant and occurs even in the cold climates of the northern boreal and subalpine. Adults will forage far from open water. Adults hibernate on land under debris and litter and rely on deep snows for insulation. The two introduced frog species, Bullfrog and Green Frog, occur on the south Coast and in the Southern Interior. These species are both highly aquatic and rarely leave natal marshes and shallow-water habitats. Stream-breeding Amphibians Paciﬁc Giant Salamander occur only in the mainland areas of the Geor- gia Depression south of the Fraser River. They breed in clear-ﬂowing streams and the adults forage in the upland forest not far from this habitat. Tailed Frog occur in and along fast-moving streams at higher elevations of the mainland Coast. Adults use adjacent upland forest and do not favour wetlands or related ecosystems. Terrestrial Amphibians Three species of lungless salamander (Plethodontidae) occur in British Columbia that live completely terrestrial lives: Western Red-back Salamander, Ensatina, and Clouded Salamander. They all occur on Vancouver Island and the adjacent mainland, except the latter, which is absent from the mainland.

222 Wetlands of British Columbia: A Field Guide to Identification These salamanders are not associated with open-water habitat even for breeding. As their common name suggests, Plethodontid salamanders have no lungs but breath through their skin and oral cavity. For this reason, they require habitats that allow them to keep their skins moist. These habitats are often moist forests with structures such as downed wood, rocks, or abundant litter under which they can hide from desiccating conditions and predators. Eggs are laid in large pieces of decayed wood that remain moist. Swamp forests are good habitat. Waterbird Habitat Requirements Wetlands provide valuable habitat for many species of birds, but waterbirds (pelicans, loons, grebes, phalaropes, swans, geese, ducks, herons, cranes, rails, shorebirds, terns) are the most dependent on wetland habitats. Natural wetlands that have seasonal and long-term fluctuations in water levels are generally the most productive for waterbirds. These fluctuations enhance productivity and maintain complex vegetation structure. Water depth and vegetation structure are important cues for waterbird use of wetland habitats (Reid 1993). Wetlands are used for different life history requirements: • summer breeding and feeding sites • moulting sites (ducks and geese) • feeding away from nesting sites (pelicans, herons, and waders) • migration stops (shorebirds, ducks, and geese) • food exploration after breeding (waders) • wintering Loons and Grebes Loons and Grebes require large stretches of open water to achieve flight and therefore occur on larger waterbodies. They nest mostly on mounds of floating vegetation (often Muskrat haul-outs) within the cover of emergent vegetation. Pacific Loons will select large lakes that are more clear than coloured, because they are sight feeders and colour may inhibit foraging efficiency. Horned Grebes are likely to occur on larger wetlands with high pH and chlorophyll levels. Eared Grebes colonies nest on surface mats of filamentous green algae, sago pondweed, or bladderwort anchored in sedges and other emergents (Boe 1994).

Chapter 6 Conservation and management issues 223 The floating nests of Red-necked Grebes are found in shallow waters of lakes.

Herons, Cranes, and Rails Great Blue Herons move from coastal, intertidal habitats during the breeding season to marshes and grasslands during November to Febru- ary (Butler 1995). This seasonal shift in habitat use occurs because herons can no longer meet their daily energy needs by foraging on beaches. Herons nest in colonies, commonly forming rookeries in red alder or cottonwood. Sandhill Cranes breed in bogs with a shrub layer, swampy ground with pink spirea, wetlands dominated by sedges, and coniferous forest (Coop- er 1996). In British Columbia, Sandhill Cranes utilize wetlands in the

Great Blue Herons are commonly seen in marshes and estuaries but nest colonially in upland forests.

224 Wetlands of British Columbia: A Field Guide to Identification 1–100 ha size range. Adults build nests out of mounds of vegetation in shallow wetlands within stands of emergent vegetation. Young and adults use coniferous forests for escape cover and possibly for resting and feeding. During migration stopovers, cranes use shallow, open wetlands with clear water for loafing, drinking, and roosting. American Bitterns prefer vegetated edges and shorelines of wetlands that are dominated by tall, emergent vegetation (i.e., taller than the bird itself). They have been found in wetlands of all sizes, but appear to prefer impoundments or beaver-created wetlands (Gibbs et al. 1992). Use of wetlands appears to be restricted to those portions with water that are shallow enough to stand in (less than 10 cm). Nests are well-concealed platforms of reeds and other vegetation within dense emergent vegetation such as cattail and bulrush. Bitterns will occasionally nest on dry ground in dense vegetation greater than 30 cm tall in grasslands adjacent to wetlands (Gibbs et al. 1992). Breeding Virginia Rails are rare to locally common in the Georgia Depression, Southern Interior, and Central Interior at low elevations. They favour regions with warm spring air temperatures. Virginia Rails use freshwater wetlands, but have also been found in salt marshes, favouring stands of robust vegetation including cattails and bulrushes (Conway 1995). They have been frequently observed in areas with 5–15 cm deep water, and heard from areas with less than 5 cm water. Nests are loosely woven baskets usually concealed with an overhead canopy. Nests can be well above the water, or at the base of taller vegetation (Harrison 1978). Soras are fairly common along southeastern Vancouver Island, Fraser Lowlands, and in suitable habitat throughout the Interior to the Peace River and other boreal areas. Soras prefer freshwater and brackish wetlands with an interspersion of emergent vegetation and open water. Cup nests are often placed at vegetation edges, near patches of open water, in a mixture of robust and fine vegetation; the surrounding vegetation is folded over to form a covering dome. Dominant plants around nest sites often include cattails and sedges, and, less commonly, bulrushes, bur- reeds, and grasses. Waterfowl Waterfowl require abundant protein and calcium during nesting and moulting. Nesting waterfowl diets require invertebrates for the protein required for egg production and growth of young. Few duck species acquire substantial nutritional resources directly from consumption of

Chapter 6 Conservation and management issues 225 plant material other than seeds. Many wetlands with apparently good vegetative cover are infrequently used by breeding or moulting waterfowl because protein-rich prey is limited. Acidic wetlands are usually poor breeding habitat because the macroinvertebrate communities are dominated by dipterans with low protein and calcium content. Saline/alkaline waterbodies in warmer climates typically have the highest density of breeding waterfowl in British Columbia. Saline lakes provide high quality and abundance of prey but broods must be moved to freshwater seeps or adjacent freshwater marshes (ducklings cannot tolerate salt levels greater than 15 ppt). Waterfowl, grebes, loons, and coots undergo a complete, simultaneous wing moult that leaves them flightless. The moult occurs during the brood-rearing period of geese, the post-breeding period of male ducks, and the post–brood-rearing period of most female ducks. Feather re- growth requires 3–5 weeks, depending on the species. Birds often move to areas away from their breeding grounds to moult. Although little research has been done, it is believed that these moulting areas are rich in food and/or offer good cover for protection from predators. Migrating Tundra Swans prefer wetlands with fennel-leaved pondweed (Stuckenia pectinata), while non-foraging swans prefer wetlands with large, open, unvegetated mud bars (Earnst 1994). Trumpeter Swans feed on plant foods almost exclusively, although the type of plants consumed by each sex varies during the season. Before in- cubating, adults feed on submerged aquatic macrophytes, while during incubation, horsetails (Equisetum fluviatile and E. arvense) and the sedge, Carex lyngbyei, become the food item of choice for females, while males continue to feed on submerged aquatics. Adult swans and their young spend more time feeding on horsetail than all other foods combined. (Grant et al. 1994). Dabbling ducks, also known as “marsh ducks,” are capable of vertical takeoff and can use small waterbodies and terrestrial habitats. Species include Mallard, Gadwall, Northern Pintail, Green-winged Teal, Blue- winged Teal, Cinnamon Teal, Northern Shoveler, American Wigeon, and Eurasian Wigeon. Distribution and breeding habitat requirements for these species are outlined in Table 6.2. Most dabbling ducks nest in a wide variety of riparian upland habitats. However, birds nesting in natural grass- and shrublands have the highest nesting success. In addition, seasonal wetlands can have higher fledging

226 Wetlands of British Columbia: A Field Guide to Identification (agricultural fields, sloughs, marshes, lakes, riparian woodlands) environments. vegetation. by concealed on ground Nests Central-southern B.C. north and Grand Forks Creston from Lake,Williams to Lowlands, the Fraser Lowlands. and Peace River and Peace abundant in Chilcotin-Cariboo areas. in the grasslandsMost and open forested Wetlands brackish Coastal parklands; Interior, throughout scattered marshes, Kootenay farmlands, sewage lagoons, and lakes. on the ground Nests Lowlands,and Nechako River, Peace and south Coast.Widespread, most abundant in the Chilcotin-Cariboo. upland habitats, in brushy and rivers wetlands Freshwater sometimes the southern east through South Coast and central Interior water. far from hanging shrubs. over- by often very on ground concealed well Nests Atlin. in urban to (golf and west forest boreal through River Wetlands and Peace courses, ditches, parks) and rural and open habitats near small bodies of Forested water. in Chicotin-Cariboo,Concentrations on Nests Okanagan, Nechako Lowlands, River. and Peace River north Powell to southern B.C.—Victoria to Confined near water. ground Lowlands; the Fraser and throughout and from Kootenays in the Lake north Nimpo to Valley southern Okanagan Wetlands, meadows, wet ditches, sewage lagoons, and slow-moving of centre Chilcotin-Cariboo—the abundance. streams. vegetation. by surrounded on ground Nests Lowlands. and Peace abundant in the Chilcotin-Cariboo Most the southernDistributed and from on the inner south Coast Open and semi-open habitats in the vicinity of marshes, sloughs, and the Peace. Atlin north Interior to ponds, the Interior,Throughout bogs, east of lakes, Ranges and locally the Coast ditches, streams. and slow-moving Lowlands. and the Queen Charlotte on the south Coast Most Drier margins of and wetlands, lakes dry grasslands,abundant in the central-south Interior, Lowlands, the Peace shrubby fields, edges ofand the east Kootenay. forests, mixed damp meadows, and subalpine bogs. Island,Vancouver Southern and Lowlands the Fraser vegetation. in sparse or low on ground Nests northern on the Coast. Lowlands Queen Charlotte Widely the Interior.distributed across marshes in the Interior; sloughs with and ponds associated estuaries Grassy, brushy, near freshwater lightly upland areas wooded the Coast. on in dense cover. on ground Nests Mallard Species Distribution habitat Breeding Gadwall Distribution and habitat use by dabbling ducks (from RIC 1999) Cinnamon Teal Northern Pintail Blue-winged Teal American Wigeon Northern Shoveler Green-winged Teal Green-winged (and Eurasian Wigeon) (and Eurasian  . 

Chapter 6 Conservation and management issues 227 success than permanent wetlands; possibly because prey quality is better than in permanent water. Optimum conditions for dabbling duck populations occur during years when large numbers of seasonal ponds contain water. However, permanent wetlands are needed to maintain adult populations during periods of drought and as moulting areas (Mauser et al. 1994). “Lake” ducks are associated primarily with lake habitats as they require sufficient open water to gain flight. “Lake” duck species in British Columbia include Redhead, Ring-necked Duck, Canvasback, Greater Scaup, Lesser Scaup, and Ruddy Duck. These ducks nest mostly in emergent vegetation of lacustrine marshes. Two sea ducks, Surf and White-winged Scoter, also nest in wetland habitats. Distribution and breeding habitat requirements for these species are outlined in Table 6.3. There are several species of cavity-nesting duck in British Columbia: Bufflehead, Common Merganser, Hooded Merganser, Common Gold- eneye, Barrow’s Goldeneye, and Wood Duck. For these species, riparian management around good-quality wetland and lake habitat is essential. Cavity-nesting ducks use Pileated Woodpecker cavities or other hollows created by decay. Bufflehead are small enough to use excavations formed by smaller woodpecker species. Distribution and breeding habitat requirements for cavity-nesting ducks are outlined in Table 6.4. Shorebirds Most shorebirds use British Columbia’s wetlands primarily during migration and wintering (Coast). They opportunistically use mudflats

A Killdeer chick hides in a coastal bog.

228 Wetlands of British Columbia: A Field Guide to Identiﬁcation Central and southern Interior to Prince George,Central Prince and southern to Interior Vanderhoof, Lowlands. the Peace and throughout Atlin and of Centre and alkali lakes, Freshwater dense emergent by bordered wetlands in Chilcotin-Cariboo.abundance Southeast Kootenays, Creston, and the Okanagan valley,widely the Cariboo-Chilcotin, throughout and in the vegetation, especially bulrush. in dense emergent on water Nests Lowlands. Peace north through the Okanagan and Creston from Widespread with and wetlands emergent vegetation. lakes freshwater On shallow and on the Lowlands the Peace to the Chilcotin-Cariboo lakes, Freshwater ofeast coast Valley. marshes, and the Fraser Island Vancouver ponds, and sloughs, in dense stands of water over in wooded often Nests emergent vegetation. northward but increasing in the southern Interior Rare and the Thompson- Trench Mountain the Rocky through situations. in grass on land or water clumps or emergent Nests and Peace the Cariboo-Chilcotin to Okanagan Plateau . areas River and alkaline lakes, in dense grass, Freshwater vegetation. concealed marshes, agricultural crops, and ﬁelds. vegetation. the east Kootenays, and Okanagan valleys to or emergent vegetation. Nicola nests Most north Lowland the Nechako to the Chilcotin-Cariboo through area. River region and in the Peace Island, as bulrushes with wetlands such emergent vegetation Freshwater Vancouver Southeastern Lowlands, the Fraser east southern B.C.,across and cattails for nesting cover. in emergent water over Nests north the Chilcotin-Cariboo through Lowlands, the Peace Basin regionsand Fraser to Fort Lowlands,Nelson Basin. and Liard Freshwater, alkali, stands of and brackish with wetlands extensive along the margins. dense emergent vegetation in on water Nests of lowlands Nelson and Fort Peace northeastern B.C.,Thompson-Okanagan stands.Atlin. to and west Lowlands the Peace to Plateau Fraser dense stands of emergent vegetation. or mature and muskeg spruce by surrounded lakes Freshwater vegetation. and ponds in open country forest. lakes and boreal Freshwater coniferous/deciduous. Species Distribution habitat Breeding Redhead Distribution and habitat use by “lake” ducks (from RIC 1999) Surf Scoter Canvasback Ruddy Duck Lesser Scaup Lesser American Coot Ring-necked Duck White-winged Scoter White-winged  . 

Chapter 6 Conservation and management issues 229 pine forests, farmland, and rangeland. near cavities in tree Nests the edge of wetlands. wetter, forests, coniferous closed including subalpine regions. preferred. are Alkaline lakes of in cavities Nests near the trees edge of a wetland. or crevices. in small caves Southern Vancouver Island,Vancouver Southern Lowlands, Fraser southernInterior, especially Creston, Lake.Williams north to Interior,Across and northern Lowlands especially the Peace ofareas forest. the boreal ponds, lowland to adjacent deciduous woodlands Mature sloughs, on rivers, and occasionally Primarily on lakes sloughs, and ponds but widespread of in the southern third Uncommon B.C. rivers. and slow-moving of in cavities Nests deciduous trees. east of mature Ranges. the Coast the north. Sparse through east of in the southern Interior Widespread the Coast Ranges. region. abundant in Chilcotin-Cariboo Most Lakes, rivers, floodplains, and associated sloughs, ponds, and with associated aspen parkland, Lakes creeks, pine of open ponderosa Centre usually with margins. wooded northern from Queen in southwest abundance cavities. in tree Nests Islands,Charlotte forests, Kitsault, farmland, St. Fort rangeland, James, and alpine meadows, and Prince as as well of the rest George south through B.C. B.C.Throughout mountainous areas. in extremely except brackish sites, but occasionally water fresh Mostly in northern and widely common scattered B.C.Less usually in aspen parklands, interior Douglas-fir forests, open ponderosa of shores along forested freshwater Near lakes, with shorelines. wooded streams, cavities. in tree Nests rivers, inlets, ponds. and beaver and on ground cavities in tree Nests Species Distribution habitat Breeding Distribution and habitat use by cavity-nesting ducks (from RIC 1999) Bufflehead Wood Duck Wood Hooded Merganser Barrow’s Goldeneye Barrow’s Common Merganser Common Common Goldeneye Common  . 

230 Wetlands of British Columbia: A Field Guide to Identification exposed during watertable drawdown. Shorebirds are attracted almost immediately to wet mud/shallow-water habitats that became available regardless of wetland history. These habitats are associated mostly with estuaries on the Coast and marshes in the Interior. Several species will nest in wetlands, including Wilson’s Phalarope, Great Yellowlegs, Lesser Yellowlegs, Spotted Sandpiper, Solitary Sandpiper, and Least Sandpiper. Other Birds’ Habitat Requirements A variety of passerines use wetlands extensively. Wetlands with high structural diversity, both vertically and horizontally, attract the most songbird species. Common passerines that occur primarily in emergent wetlands include Common Yellowthroat, Red-winged Blackbird, Marsh Wren, Swamp Sparrow, Lincoln’s Sparrow, Yellow-headed Blackbird, Le Conte’s Spar- row, and Nelson’s Sharp-tailed Sparrow. Many other species extensively use wetland-upland edge habits and riparian deciduous shrubbery for nesting and foraging. These habitats provide cover, access to open habitat, high prey density, and multilay- ered structure. Some shrub ecotone–affiliated species include the Empidonax flycatchers, Eastern Kingbird, chickadee species, Hermit Thrush, Gray Catbird, Bohemian Waxwing, Cedar Waxwing, Tennessee Warbler, Yellow Warbler, Black-and-white Warbler, American Redstart, Northern Waterthrush, MacGillivray’s Warbler, Wilson’s Warbler, Yel- low-breasted Chat, Savannah Sparrow, Fox Sparrow, Song Sparrow, Lincoln’s Sparrow, White-throated Sparrow, Black-headed Grosbeak, and Rusty Blackbird.

A Yellow-headed Blackbird in typical habitat.

Chapter 6 Conservation and management issues 231 Eastern Kingbirds commonly use dead shrubs in flooded lake margins.

There are a few passerines that frequently use bog habitats, particularly in the Boreal and Taiga Plains where these habitats are extensive. These include Ruby-crowned Kinglet, Hermit Thrush, Palm Warbler, and Blackpoll Warbler. Ruffed Grouse use shrubby riparian areas, particularly in the winter time. Willow Ptarmigan extensively use subalpine fens and shrub-carrs for forage. Raptors and Owls will use wetland habitats where abundant prey exists. Northern Harrier nest on the ground, often in the emergent cover of marshes. Bald Eagle and Osprey will nest in larger riparian cottonwood. Western Screech Owl frequently use floodplain forests. Great Gray Owl are often associated with bog habitats. Woodpecker species are not generally considered wetland users. However, forested wetlands and riparian forests are more likely to escape wildfire and become old forests. These old forests have large stems and abundant snags that are optimal for creation of nest cavities. The blue-listed Lewis’ Woodpecker nest in floodplain cottonwood adjacent to grasslands. Woodpecker excavations in riparian areas are critically important for cavity-nesting ducks. Wetland Mammal Habitat Requirements There are several mammal species that are semi-aquatic and associated primarily with open water. These include three rodents (Beaver, Muskrat, Mountain Beaver) and two mustelids (Mink and River Otter). Beaver

232 Wetlands of British Columbia: A Field Guide to Identiﬁcation and Muskrat play key ecological roles because they build structures that modify the environment and are used by many other wetlands animals. All but Mountain Beaver are also considered fur-bearers and are of eco- nomic importance. Beaver The Beaver is perhaps the most well-known wetland rodent species due to its role in wetland creation. Through dam-building, Beaver create numerous shallow-water habitats that are used extensively by many other wildlife species, especially waterfowl. Furthermore, Beaver rely heavily on riparian vegetation for forage and dam-building materials. These activities have a large impact on riparian plant communities: they cut down the dominant species (trees and shrubs), which allows understorey species to ﬂourish; cutting is concentrated in the riparian zone; and Beaver harvest in excess of their nutritional needs (Johnson and Naiman 1990). Populus tremuloides is the primary food species but Populus balsamifera, Salix spp., and Betula papyrifera are important secondary sources. Other species such as Cornus stolonifera, Amelanchier alnifolia, Corylus cornuta, and Betula nana are other minor forage species. Alnus species are used primarily for dam-building. Fire is an important disturbance agent for creating optimal forage for Beaver.

Beaver is a keystone species that creates and modifies wetland habitats.

Chapter 6 Conservation and management issues 233 Muskrat Muskrats are the most significant vertebrate consumer of emergent vegetation in North American wetlands. Muskrat foraging and house-building improves interspersion in dense stands of emergent vegetation, which increases invertebrate population levels and may result in increased avian abundance and diversity (Clark and Kroeker 1993). Muskrat haul- outs are used for nesting by species of grebe and diving duck. Cattail and bulrush in 0.5–1.5 m deep water is optimal habitat for lodge-building. Mountain Beaver Mountain Beaver is a red-listed species in British Columbia. They occur only in the southwest, coast-transition areas of the Cascade Range. They are associated with small streams or seeps in montane and subalpine forests between 1200 and 1800 m elevation where they build tunnels and dens in fine-textured soils. Primary food species often occur in wet forests or meadows and include Valerian sitchensis, Equisetum spp., Streptopus spp., Rubus parviflorus, and Heracleum maximum. Logging activities that disturb soils appear to have a strongly negative impact on Mountain Beaver populations (Gyug 2000). Mink American Mink are semi-aquatic predators that spend much of their time in water. They are a major predator of muskrat, fish, marine invertebrates, frogs, waterfowl, and eggs. High densities of mink will occur in habitats with high populations of potential prey. They occur throughout the province along fish-bearing streams and marshes. They will nest in muskrat or beaver burrows. Mink favour good riparian cover, and clearing or heavy grazing of wetland edges will reduce habitat quality. Other Mustelids River Otters use a wide range of wetland habitats throughout the year with beaver-created wetlands favoured (Newman and Griffen 1994). Marten and Fisher use large trees in riparian forest for denning. Other species such as Short-tailed Weasel, Common Raccoon, and Striped Skunk also use wetlands, particularly marshes for hunting. Small-mammal Habitat Requirements Of the 32 species of small terrestrial mammal and 16 species of bat that occur in British Columbia, only some are associated with wetlands and related ecosystems.

234 Wetlands of British Columbia: A Field Guide to Identiﬁcation Bats In a review of the importance of riparian habitat in British Columbia to bats, Brigham (1993) summarizes available information about foraging and roosting in various riparian habitats. The following species have been detected foraging over calm water: California Myotis, Western Small-footed Myotis, Long-eared Myotis, Western Long-eared Myotis, Keen’s Long-eared Myotis, Northern Long-eared Myotis, Little Brown Myotis, Yuma Myotis, Big Brown Bat, and Spotted Bat. As well, Townsend’s Big-eared Bat has been captured at streams, lakes, and small marshes dominated by willow, poplar, and birch species. All of these species feed on ﬂying insects and focus on wetland habitats with abundant insect populations. Jumping Mice Meadow Jumping Mouse are found throughout the northern half of British Columbia, east of the Coast Ranges in association with brushy margins of streams and in marshes. Western Jumping Mouse are found throughout mainland British Co- lumbia except in the extreme southwest and Taiga Plains. They use a variety of habitats but they prefer dense cover with lush grass and herb growth near open water. Voles and Lemmings Long-tailed Vole occur throughout mainland British Columbia. They use variable habitats that include marshes and alder or willow thickets along watercourses. Meadow Vole occur throughout the Interior. They use habitats with high ground cover of sedges, grasses, and forbs, usually near water. Southern Red-backed Vole are found throughout the province except the coastal islands and Boreal Mountains and Plateaus. They forage in bogs and forested swamps. Townsend’s Vole occur on Vancouver Island and the Fraser Delta in a variety of habitats, which include marshes, estuaries, and sedge meadows. Northern Bog Lemming occur throughout the province except on the islands. Although this species primarily inhabits bogs, it may also be present in other wet habitats such as deep mossy spruce woods, wet subalpine meadows, and alpine tundra.

Chapter 6 Conservation and management issues 235 Shrews Five of the 11 shrew species in British Columbia are often associated with wetlands: Arctic Shrew, Common Water Shrew, Dusky Shrew, Pacific Water Shrew, and Pygmy Shrew. These are primarily insectivores but will prey opportunistically on other small animals. Generally, they use a wide variety of habitats of which wetlands are often focal. The Arctic Shrew is restricted to the Taiga Plains where it forages in tamarack and spruce swamps, in marshy areas, and along the edge of bogs and marshes in alder or willows. Common Water Shrew occur throughout the province (except the Queen Charlotte Islands) usually in or near water. Lakes, ponds, swift and sluggish small streams, cold fast mountain streams, and stagnant water of bogs and marshes are all used. Dusky Shrew are widespread at all elevations. They use a wide variety of different habitats under a wide range of climatic conditions but are frequently found in marshes, streamside Equisetum stands, and bogs. The red-listed Pacific Water Shrew has a limited distribution in British Columbia, occurring only in the Fraser River delta where it uses a wide variety of habitats but favours riparian red alder forests or forested skunk cabbage swamps. Pygmy Shrew are found throughout the Interior and are generally uncommon. They use a wide variety of habitats, which include bogs and marshes. Ungulate Use of Wetlands and Related Habitats Moose are the most wetland-associated ungulate in British Columbia. Browse makes up 80% of the diet, and favoured species such as willow and red-osier dogwood are associated with wetlands. An overview of the forage value of 15 willow species found in the Lower Skeena and Bulkley valleys is presented by Roberts (1986). Moose also use aquatics in the summer and will dive for Nuphar rhizomes. Elk dwell primarily in open forest habitats in dry climates. They are not specifically wetland users but will graze sedges where they are available. Mule Deer and White-tailed Deer will use wetland habitats for cover, especially in otherwise open terrain.

236 Wetlands of British Columbia: A Field Guide to Identiﬁcation Coastal estuaries are high-quality habitat for many species, including Black-tailed Deer.

Bear Use of Wetlands and Related Habitats Some wetlands are important Grizzly Bear and Black Bear habitats. Productive swamp and estuarine habitats typically support forbs and fruiting shrubs favoured by bear. Important spring forage species include (bold are major forage species): Athyrium filix-femina, Calamagrostis canadensis, Carex lyngbyei, Conioselinum pacificum, Empetrum nigrum, Equisetum arvense, Heracleum maximum, Lysichiton americanus, Oplopanax horridus, Populus balsamifera, Scirpus microcarpus, and Taraxacum officinale. In summer and fall, the fruits of many shrubs and forbs are consumed: Arctostaphylos uva-ursi, Cornus stolonifera, Corylus cornuta, Empetrum nigrum, Gaultheria shallon, Lonicera involucrata, Malus fusca, Oplo- panax horridus, Ribes bracteosum, Rosa spp., Rubus idaeus, R. parviflorus, R. spectabilis, Sambucus racemosa, Streptopus spp., Vaccinium spp., Valeriana sitchensis, and Viburnum edule. Macroinvertebrates Macroinvertebrates, including annelid worms, molluscs, insects, and crustaceans, are extremely important components of wetland ecosystems. They convert plant matter to high-protein prey that support many wetland vertebrates. The highest macroinvertebrate populations are frequently associated with wetlands with high levels of annual vegetation growth. Sites with abundant vegetation growth have good stuctural complexity important

Chapter 6 Conservation and management issues 237 for many life history requirements: forage, escape cover, sites for ovipo- sition, emergence, respiration, attachment, and pupation. The most productive wetlands are those with fluctuating watertables, and non-acid waters (marshes). Drawdowns facilitate nutrient release and foster plant growth that contributes to nutrient and detritus supply on reflooding. Sites with permanent and stable water levels have lower macroinvertebrate biomass because litter quantities are reduced, but they frequently have more diverse communities with a full range of detritivores and their predators. Inland saline waters are often highly productive for waterfowl, in part because these habitats lack fish competitors for prey. Predation by fish is a profound influence on invertebrate abundance and community structure and abundance. Coleopterans, Hemipterans, and Amphipods are common in saline waters.

A Swallowtail on the rare Henderson's checker-mallow (Sidalcea hendersonii).

238 Wetlands of British Columbia: A Field Guide to Identiﬁcation APPENDIX 1 Ordination of Site Associations

The following ordination diagrams are based on average species cover values for the Site Associations described in this guide. In general, ecosystems that occur close together in the diagrams are similar in species composition. The axes of the diagrams are not directly related to environmental gradients, but correlations to gradients can be inferred from known attributes of the ecosystems ordinated. Ordinations were generated by Nonmetric Multidimensional Scaling () for average species cover values for the Site Associations described in this guide. Site Associations are identified by letter code (or a short name where units have not been coded) and also symbol-coded by Site Class. All ordinations were performed using the  autopilot routine of - 4.0 (McCune and Mefford 1999) with the following attributes: Distance measure = SORENSEN, Number of axes = 4, Ran- dom number seed starting configuration, 15 runs with real data and 30 runs with randomized data. Figure 1.1 is an ordination of all numbered Site Associations used in the guide including some representative shallow-water ecosystems. Final stress for 3-dimensional solution was 14.74 (Monte Carlo p<0.0323) reached after 166 iterations with a final instability of 0.00010. Figure 1.2 is an ordination of only numbered wetland Site Associations. Final stress for 3-dimensional solution was 12.23 (Monte Carlo p<0.0323) reached after 91 iterations with a final instability of 0.00008.

Appendix 1 239 Dry Class Bog Ws53 Em03 Fen Fl51 Marsh Swamp, shrub Swamp, forest Fm50 Shallow-waters Fm03 Estuarine Ws51 Low Bench Fl02 Ws54 Middle Bench Ws11 Fl50 Fl01 Fm01 Meadow Ws55 Fm02 Ws10 Fl03 Fl06 Shrub-carr Fl04 Ws01 Ws08 Wm50 Em07 Ws09 Ws07 Fl07 Ws06Ws03 Wb53 Ed02 Wf52 Fl05 Ed01 Wb52Wb03Wb09 Ws02 Wb04 Ws50 Em06 Em04 Wb07 Wb51 Wb01 Wb08 Sc03 Gs03 Wf51 Wb05 Em05 Wb02 Wb11Wf50 Wf04 Wb50 Wb06 Wf03 Ws04 Schoame Wb10 Wf02 Wf12 Wb13 Wf13 Wf53 Ws05 GS04 Gs01 Wf07 Wf11 Sc02 Sc01 Wm07 Wb12 Wm05 Dynamic Bolbmar Wm51 Wf09Wf08 Wf05 Wf01Wm01

Wm03 Wf10 Wf06 Wm06 Gs02 Wm02 Nuphlut Em01 Hydrodynamic Trigmar Wm04 index

Stagnant Brassch

Wet Potamog

Poor Rich Saline

 . NMS ordination of numbered wetland and related Site Associations including some shallow-water units.

240 Wetlands of British Columbia: A Field Guide to Identiﬁcation Dry Ws53 Class Bog Fen Marsh Swamp, shrub Swamp, forest

Ws51

Ws54

Ws11 Ws55 Ws01 Wm50 Ws10

Ws06 Ws07 Ws50 Ws03 Ws09 Wf52 Ws02 Ws08 Wb53 Wb04 Wf50 Wm07 Wb03 Wb09 Wb07 Wf04 Wb52 Ws04 Wf51 Wb08Wf03 Wb50 Wb05 Wb51 Wb01 Wb11 Wf53 Wb02 Wm05 Wb06 Wf02 Wb10 Ws05 Wm51 Wf01Wm01 Dynamic Acid Wb13 Wf12 Wf13 Wm03 Wf07 Wb12 Wf11 Wf05 Wf09 Wm02 Wf08 Hydrodynamic index pH Wm06 Wf06 Wm04

Stagnant Wf10 Wet Neutral

Poor Rich

 . NMS ordination of numbered wetland Site Associations only.

Appendix 1 241 APPENDIX 2 Overlays of the Wetland Edatopic Grid

The following overlays on the wetland edatopic grid indicate the ecological space of various vegetation characteristics. Woody plants are limited by excessive moisture or prolonged flooding. Figure 2.1 indicates the approximate areas of the grid where physiognomic types are likely to occur. Forest occurs on very moist sites or wet sites where there are sufficient elevated microsites to support good tree growth. Trees and shrubs <10 m in height can occur on wet sites but wetter sites will not support tall shrub communities. Some very wet sites will support low shrubs where sufficient microtopography exists.

Soil Nutrient Regime ABCD E F

May support forest VM

W May support tall shrub AM ANAkVA N MA SA May support low shrub VW Soil Moisture Regime Herb or Bryoid pH

St Sl Mo Dy VD Hydrodynamic Index

 . Edatopic grid position of vegetation physiognomy.

Figure 2.2 shows the grid location of peat-accumulating ecosystems (peatlands) and dominant bryophyte groups or species. Stagnant and sluggish sites are peatlands and are accumulating peat. Sites with mobile waters often have some thin, well decomposed peat veneers, while more dynamic sites will have minimal surface organic matter. Sphagnum mosses dominate in acidic peat-accumulating sites, while more minerotrophic species occur in slightly acidic to alkali peatlands. Wetlands with more dynamic hydrology often have low bryophyte cover because of prolonged deep surface flooding or excessive shading and litterfall from a vigorous deciduous canopy. Broad classes from vegetation classification are defined based on vegetation similarity, which does not exactly match site class boundaries. Eight

242 Wetlands of British Columbia: A Field Guide to Identiﬁcation Soil Nutrient Regime ABCD E F Feathermosses M Hylocomium splendens Rhytiadelphus Pleurozium VM

Sphagnum Aulacomium W Group I & III Mnium spp. Aulocomium AM ANAkVA N MA SA Few mosses Brown mosses T. nitens, S. Group II VW Drepanocladus Soil Moisture Regime S. warnstorfii pH Campylium Scorpidium

Peat-accumulating St Sl Mo Dy VD sites Hydrodynamic Index

 . Edatopic grid position of major bryophyte groups and peat-accumulating sites. vegetation classes that cover the range of wetlands and related ecosystems are presented in Figure 2.3. Meidinger et al. (2001) provide full descriptions of the following wetland and related vegetation classes. Class Oxycoccus oxycoccos – Sphagnum Communities of interior and coastal bogs and poor fens with stagnant, acidic, base-poor, organic soils. Characterized by the prevalence of Sphagnum species in the bryophyte layer and of other species tolerant of saturated, highly acidic soils. Site conditions are wet or very wet and nutrient-poor to very poor. Orders of this Class reflect climatic variation. Class Populus balsamifera – Alnus Forests throughout province dominated by, or with a significant component of, Populus balsamifera, or (for floodplains) associated with Populus balsamifera forests. Class Carex – Drepanocladus Communities of interior fens with high, stable watertables and high base-cation availability. Variously dominated by Cyperids such as Carex lasiocarpa, C. limosa, Eleocharis quinqueflora, Trichophorum cespitosum, or T. alpinum and with brown mosses such as Drepanocladus species, Campylium stellatum, Scorpidium scorpioides, or Tomentypnum nitens. Class Magnocarex Communities of interior peatlands and flooded mineral substrates dominated by large sedges, especially Carex aquatilis, C. sitchensis, or C. utriculata, but also other large cyperids such as C. atherodes,

Appendix 2 243 Soil Nutrient Regime ABCD E F Class M Hordeum jubatum Coniferous Class or Classes Populus Class VM balsamifera – Carex Alnus lyngbyei

W Class Oxycoccus oxycoccus – Class Typha – AM ANAkVA N MA SA Sphagnum Class Schoenopectus Magnocarex – Eleocharis

VW Soil Moisture Regime Class pH Carex limosa – Drepanocladus

St Sl Mo Dy VD Hydrodynamic Index

 . Edatopic grid position of Classes from the British Columbia Vegetation Classification (Meidinger et al. 2001).

C. exsiccata, Dulichium arundinaceum, or Scirpus microcarpus. Site conditions are Wet to Very Wet with a circum-medium nutrient regime and slightly acidic to circum-neutral soils. Shallow flooding followed by growing-season drawdown to just below the surface is typical. These are tall shrub, low shrub, or graminoid-dominated communities with large Cyperaceae always being prominent. Class Typha – Schoenoplectus – Eleocharis Communities of interior and coastal marshes deeply flooded with mobile, often strongly fluctuating watertables, and abundant available nutrients. Often simple communities strongly dominated by one to few emergent grass or grass-like species. Class Hordeum jubatum Interior communities of saline or alkali meadow/marshes occurring in warm and semi-arid climates. Located in basins and lake margins where evaporative drawdown accumulates mineral precipitates. These are low- diversity communities with flood- and salt- or alkali-tolerant grasses and forbs such as Carex praegracilis, Distichlis spicata var. stricta, Hordeum jubatum, Juncus balticus, Poa juncifolia, Puccinellia nuttalliana, Salicornia rubra, Suaeda depressa, Spartina gracilis, and Triglochin maritima.

244 Wetlands of British Columbia: A Field Guide to Identiﬁcation APPENDIX 3 Wetland Indicator Plant Species

The following tables includes all plant species used in this guide and their value as indicators of wetland environments. The water-saturated environment of wetlands supports a unique group of plants called “hydrophytes.” These plants are adapted to grow in waterlogged soils. Species are designated as one of three indicator types. Obligate Hydrophytes – Species that occur only under wetland conditions (Table 3.1). Facultative – wetland afﬁliated – Species that grow primarily under wetland conditions but also, less commonly, in uplands (Table 3.2). Facultative – upland afﬁliated – Species that occur in wetlands but are widespread in uplands (Table 3.3).

 . List of Obligate Hydrophytes Shrubs Chamaedaphne calyculata Salix athabascensis Salix candida Salix pedicellaris

Herbs and Dwarf Shrubs Bolboschoenus fluviatilis Carex utriculata Oxycoccus oxycoccos Bolboschoenus maritimus Carex viridula Phragmites australis Carex aquatilis Cicuta bulbifera Polygonum amphibium Carex atherodes Comarum palustre Ranunculus gmelinii Carex buxbaumii Drosera anglica Ranunculus sceleratus Carex canescens Drosera rotundifolia Rhynchospora alba Carex chordorrhiza Dulichium arundinaceum Rubus arcticus ssp. acaulis Carex cusickii Eleocharis palustris Rubus chamaemorus Carex diandra Eleocharis quinqueflora Sanguisorba officinalis Carex exsiccata Equisetum fluviatile Sarracenia purpurea Carex flava Eriophorum angustifolium Scheuchzeria palustris Carex gynocrates Eriophorum brachyan- Schoenoplectus acutus Carex lanuginosa therum Schoenoplectus tabernae- Carex lasiocarpa Eriophorum chamissonis montani Carex leptalea Glyceria borealis Scirpus microcarpus Carex limosa Glyceria grandis Scutellaria galericulata Carex livida Glyceria occidentalis Sium suave Carex lyngbyei Hypericum anagalloides Trichophorum alpinum Carex magellanica Iris pseudacorus Trichophorum cespitosum Carex pauciflora Kalmia microphylla Triglochin maritima Carex pluriflora Lycopodiella inundata Triglochin palustris Carex sitchensis Lycopus uniflorus Typha latifolia Carex tenuiflora Menyanthes trifoliata Viola palustris

Note: Species in bold can be very abundant or dominant on wetland sites.

Appendix 3 245  . Continued

Aquatics Brasenia schreberi Nuphar lutea Sparganium emersum Calla palustris Potamogeton natans Sparganium eurycarpum Ceratophyllum demersum Potamogeton richardsonii Spirodela polyrhiza Chara spp. Ranunculus aquatilis Subularia aquatica Elodea canadensis Ricciocarpos natans Utricularia intermedia Hippuris vulgaris Ruppia maritima Utricularia macrorhiza Lemna minor Schoenoplectus subterm- Vallisneria americana Lemna trisulca inalis Zostera marina Myriophyllum verticillatum Sparganium angustifolium

Lichens and Mosses Calliergon cordifolium Scorpidium scorpioides Sphagnum pacificum Calliergon giganteum Siphula ceratites Sphagnum palustre Calliergon richardsonii Sphagnum angustifolium Sphagnum papillosum Calliergon stramineum Sphagnum austinii Sphagnum rubellum Calliergonella cuspidata Sphagnum balticum Sphagnum russowii Campylium stellatum Sphagnum capillifolium Sphagnum subnitens Drepanocladus sendtneri Sphagnum centrale Sphagnum subsecundum Fontinalis antipyretica Sphagnum compactum Sphagnum tenellum Hamatocaulis lapponicus Sphagnum contortum Sphagnum teres Hamatocaulis vernicosus Sphagnum cuspidatum Sphagnum warnstorfii Hypnum lindbergii Sphagnum fimbriatum Tomentypnum nitens Meesia longiseta Sphagnum fuscum Warnstorfia exannulata Meesia triquetra Sphagnum lindbergii Warnstorfia fluitans Paludella squarrosa Sphagnum magellanicum Warnstorfia sarmentosa Philonotis fontana Sphagnum majus Scorpidium revolvens Sphagnum mendocinum

 . List of Facultative Hydrophytes – Wetland Affiliated

Trees Larix laricina Pinus contorta var. contorta

Shrubs Alnus incana Salix barrattiana Betula occidentalis Salix lucida Ledum groenlandicum Salix maccalliana Myrica gale Spiraea douglasii

246 Wetlands of British Columbia: A Field Guide to Identiﬁcation  .. Continued

Herbs and Dwarf Shrubs Agrostis aequivalvis Equisetum palustre Petasites frigidus Agrostis scabra Equisetum sylvaticum Petasites sagittatus Alopecurus aequalis Equisetum variegatum Pinguicula vulgaris Amphiscirpus nevadensis Fauria crista-galli Plantago maritima Anagallis minima Galium trifidum Platanthera dilatata Anemone occidentalis Gaultheria hispidula Platanthera stricta Angelica arguta Gentiana douglasiana Puccinellia nutkaensis Angelica genuflexa Gentiana sceptrum Puccinellia nuttalliana Aster borealis Glaux maritima Puccinellia pumila Aster modestus Glyceria elata Ranunculus aquatilis var. Beckmannia syzigachne Glyceria striata diffusus Berula erecta Juncus arcticus Ranunculus orthorhynchus Bolboschoenus maritimus Juncus balticus Rorippa palustris Calamagrostis stricta Juncus effusus Rubus arcticus Caltha leptosepala Juncus ensifolius Salicornia virginica Carex anthoxanthea Kobresia sibirica Sanguisorba canadensis Carex capillaris Leptarrhena pyrolifolia Sanguisorba menziesii Carex disperma Lilaea scilloides Schoenoplectus americanus Carex echinata Lobelia kalmii Schoenoplectus saximon- Carex nigricans Lysichiton americanus tanus Carex obnupta Lythrum salicaria Scolochloa festucacea Carex stylosa Maianthemum trifolium Senecio atropurpureus Cicuta douglasii Microseris borealis Spartina gracilis Coptis aspleniifolia Mimulus guttatus Spartina patens Coptis trifolia Muhlenbergia glomerata Torreyochloa pauciflora Distichlis spicata Parnassia fimbriata Triantha glutinosa Dodecatheon jeffreyi Parnassia palustris Trientalis europaea ssp. Epilobium palustre Pedicularis parviflora arctica

Lichens and Mosses Aulacomnium palustre Mylia anomala Rhizomnium pseudopunc- Brachythecium salebrosum Philonotis fontana tatum Brachythecium turgidum Plagiomnium ellipticum Sphagnum squarrosum Campylopus atrovirens Pohlia nutans Drepanocladus aduncus Pohlia wahlenbergii

Appendix 3 247  . List of Facultative Hydrophytes – Upland Affiliated

Trees Abies amabilis Physocarpus capitatus Picea sitchensis Abies lasiocarpa Picea engelmannii Pinus contorta var. latifolia Alnus rubra Picea engelmannii x glauca Thuja plicata Betula papyrifera Picea glauca Tsuga heterophylla Malus fusca Picea mariana Tsuga mertensiana

Shrubs Betula nana Ribes bracteosum Salix drummondiana Cornus stolonifera Ribes lacustre Salix glauca Crataegus douglasii Rosa acicularis Salix pseudomonticola Elliottia pyroliﬂorus Rubus spectabilis Sambucus racemosa Juniperus communis Salix alaxensis Vaccinium alaskaense Ledum glandulosum Salix barclayi Vaccinium ovalifolium Lonicera involucrata Salix bebbiana Vaccinium uliginosum Menziesia ferruginea Salix brachycarpa Viburnum edule Pentaphylloides ﬂoribunda Salix commutata

Herbs and Dwarf Shrubs Achillea millefolium Epilobium ciliatum Phalaris arundinacea Agrostis exarata Equisetum arvense Plantago macrocarpa Aster subspicatus Equisetum pratense Poa palustris Athyrium filix-femina Erigeron peregrinus Poa pratensis Blechnum spicant Fragaria virginiana Potentilla anserina Calamagrostis canadensis Gaultheria shallon Pyrola asarifolia Calamagrostis nutkaensis Geum macrophyllum Pyrola chlorantha Carex praegracilis Gymnocarpium dryopteris Pyrola minor Carex scoparia Heracleum maximum Rubus parviflorus Claytonia cordifolia Hordeum brachyantherum Rubus pedatus Cornus canadensis Hordeum jubatum Salicornia maritima Cornus suecica Juncus arcticus ssp. sitch- Senecio triangularis Cotula coronopifolia ensis Spergularia canadensis Danthonia intermedia Lilaeopsis occidentalis Streptopus amplexifolius Delphinium glaucum Lycopodium annotinum Suaeda calceoliformis Deschampsia cespitosa Maianthemum dilatatum Trollius albiflorus Distichlis spicata var. Maianthemum stellatum Urtica dioica stricta Matteuccia struthiopteris Vaccinium vitis-idaea Dryopteris expansa Mentha arvensis Vahlodea atropurpurea Dryopteris filix-mas Mitella nuda Valeriana sitchensis Empetrum nigrum Mitella pentandra Veratrum viride Epilobium angustifolium Muhlenbergia richardsonis

Lichens and Mosses Cladina rangiferina Pellia neesiana Rhytidiadelphus loreus Cladonia borealis Pleurozium schreberi Rhytidiadelphus triquetrus Hylocomium splendens Polytrichum juniperinum Sphagnum girgensohnii

248 Wetlands of British Columbia: A Field Guide to Identiﬁcation APPENDIX 4 Crosswalk to other classifications

 . Wetland and related ecosystem Site Associations with corresponding BEC site series

BEC Site Series

Fl02 ICHvc/52, ICHwc/52 Fl50 CWHdm/10, CWHds1/10, CWHds2/10, CWHmm1/10, CWHms1/09, CWHms2/09, CWHvm1/11, CWHwm/07, CWHws1/09, CWHws2/09, CWHxm/10 Fl51 CWHvh1/10, CWHvh2/10, CWHwh1/09 Fm01 BGxw1/08, IDFdk1a/94, IDFxh1a/98, IDFxh2a/95, PPdh2/04 Fm02 ICHmc1/05, ICHmc2/06, SBSdk/08, BWBSmw2/05 Fm03 ICHvc/05, ICHwc/06 Fm50 CDFmm/08, CWHdm/09, CWHds1/09, CWHds2/09, CWHmm1/09, CWHms1/08, CWHms2/08, CWHvm1/10, CWHvm2/10, CWHws2/08, CWHxm/09 Wb01 SBSdk/09 Wb02 ICHmm/07, SBSvk/08, SBSwk3/05, SBSwk3a/05 Wb03 BWBSmw2/08, BWBSdk1/10, BWBSdk2/07, BWBSmw1/08 Wb05 ICHmc2/08, SBPSdc/07, SBPSmc/07, SBPSmk/08, SBSdh1/08, SBSdh2/08, SBSmc2/12, SBSmc3/09, SBSmk1/10, SBSmw/10, SBSwk1/11 Wb06 BWBSdk2/08 Wb08 SBPSdc/07, SBPSmk/08, SBSdh/08, SBSmk1/10, SBSdk/10, SBSdw2/11, SBSdw3/10 (in part) Wb09 BWBSdk1/09, BWBSwk1/07, BWBSwk2/07 Wb51 CWHvh2/31 Wb52 CWHvh2/32 Wb53 CWHvh2/12 Wf01 ICHmk1/08 Wf03 ESSFdc1/07, ESSFdc2/09, ESSFvc/06, ESSFwc1/05, ESSFwc2/10 Ws01 ICHvc/52, ICHwc/52 Ws07 ICHdk/09, ICHmc2/07, ICHmk2/06, ICHvc/06, ICHwc/08, IDFdk2/06, IDFdk3/09, IDFdk4/10, IDFdm2/07, IDFxm/09, MSdc2/08, MSdk/06, MSdm2/07, MSdv/09, MSxv/08, SBPSdc/08, SBPSmc/06, SBPSmk/07, SBSdk/07, SBSdw1/09, SBSdw3/09, SBSmc1/08, SBSmc2/10b, SBSmc2/11, SBSmc3/08, SBSmk1/09, SBSmk2/06, SBSmw/09, SBSwk1/09, SBSwk2/06 Ws08 ESSFdc1/06, ESSFdk/06, ESSFdv/06, ESSFmc/09, ESSFmc/10, ESSFmk/06, ESSFmm1/07, ESSFmv1/05, ESSFmv2/06, ESSFmv3/07, ESSFmv4/05, ESSFmw/08, ESSFvc/05, ESSFwc3/03, ESSFwc4/07, ESSFwk1/07, ESSFwv/09, ESSFxc/08 Ws10 ICHmw2/08, ICHmw3/08, ICHvk1/06, ICHvk2/06, ICHwk1/08, ICHwk2/08, ICHwk3/09 Ws11 SBSvk/10 Ws53 CDFmm/11, CWHxm/12 Ws54 CWHds1/12, CWHds2/12, CWHmm1/12, CWHms1/11, CWHms2/11, CWHvh1/13, CWHvh2/13, CWHvm1/14, CWHwh1/12, CWHwh2/06, CWHws1/11, CWHws2/11 Ws55 MHmm2/09, MHwh1/09, MHwh2/09

Appendix 4 249  . Wetland Site Associations corresponding to the Cariboo wetland classification

Cariboo IDF (Steen and Roberts 1988) Cariboo SBPS (Roberts 1984) Gs01 Alkali saltgrass – Nuttall’s alkaligrass Saltgrass – Alkaligrass meadow meadow (IDFb2/W1) (SBSa/W1) Gs02 Nuttall’s alkaligrass – Foxtail barley meadow (IDFb2/W2) Gs03 Arctic rush – Field sedge meadow Arctic rush – Field sedge meadow (IDFb2/W3) (SBSa/W2) Gs04 Tufted hairgrass meadow (SBSa/W3) Sc01 Scrub birch – Kinnikinnick shrub-carr Scrub birch – Kinnikinnick shrub-carr (IDFb2/W4) (SBSa/W4) Sc02 Grey-leaved willow – Moss shrub-carr (SBSa/W5) Wb01 Labrador tea – Sphagnum bog SBSa/W9) Wf01 Beaked sedge – Water sedge fen Beaked sedge fen (SBSa/W12) (IDFb2/W9) Wf05 Slender sedge – Moss fen (IDFb2/W10) Slender sedge fen (SBSa/W14) Wf06 Buckbean – Slender sedge fen (IDFb2/W11) Wf07 Low willow – Buckbean fen Bog willow – Sedge low shrub fen (IDFb2/W7) (SBSa/W8) Wm03 Awned sedge fen-marsh (IDFb2/W8) Awned sedge marsh (SBSa/W11) Wm05 Cattail marsh (IDFb2/W13) Wm06 Great bulrush marsh (IDFb2/W14) Bulrush marsh (SBSa/W15) Wm07 Arctic rush – Field sedge meadow Arctic rush – Field sedge meadow (IDFb2/W3) (SBSa/W2) Ws04 Drummond’s willow – Sedge swamp (SBSa/W7) Ws05 MacCalla’s willow – Beaked sedge fen MacCalla’s willow – Sedge tall shrub fen (IDFb2/W6) (SBSa/W6)

250 Wetlands of British Columbia: A Field Guide to Identiﬁcation  . Wetland Site Associations corresponding with Klinka et al. 1997

Fl04 Equisetum arvense – Salix sitchensis (in part) Fl50 Equisetum arvense – Salix sitchensis (in part) Wb50 Ledum groenlandicum – Sphagnum Assoc. Wf50 Eriophorum – Carex aquatilis Assoc. Wf52 Carex lasiocarpa and sitchensis – Myrica gale Assoc. Wf53 Nuphar polysepala: Carex lasiocarpa Subassoc. Wm04 Nuphar polysepala: Eleocharis paustris Subassoc. Wm05 Juncus ensifolius – Typha latifolia Assoc., Oenanthe sarmentosa – Typha latifolia Assoc. Wm06 Nuphar polysepala: Scirpus validus Subassoc. Wm51 Menyanthes trifoliata – Dulichium arundinaceum Assoc. Ws50 Spiraea douglasii – Carex sitchensis, Spiraea douglasii – Carex phyllomanica and sitchensis Assoc. Ws51 Lysichiton americanum – Salix lasiandra and sitchensis Assoc. Ws52 Carex obnupta – Alnus rubra Assoc.

Appendix 4 251  . Wetland Site Associations corresponding with coastal classifications in Washington and Alaska

Coastal Washington (Kunze 1994) Copper River Delta, Alaska (Boggs 2000) Em04 Puccinellia nutkaensis c.t. Em05 Carex lyngbyei c.t. Carex lyngbyaei c.t. Em06 Carex lyngbyaei – Mixed-herb c.t. Fl01 Alnus incana c.t.* Fl04 Salix sitchensis c.t. Fl50 Cornus stolonifera – Salix sitchensis c.t. Salix sitchensis c.t. Fl51 Alnus rubra/Rubus spectabilis c.t. Wb50 Kalmia occidentalis – Ledum groenlandicum/Sphagnum spp. c.t. Wf04 Salix barclayi – Carex sitchensis c.t. Wf50 Wf51 Carex sitchensis – Sphagnum c.t. Carex sitchensis – Sphagnum c.t. Wf52 Myrica gale c.t. Wf53 Carex lasiocarpa c.t. Wm01 Carex rostrata c.t. Carex rostrata c.t. Wm02 Equisetum ﬂuviatile c.t. Equisetum ﬂuviatile c.t. Wm04 Eleocharis palustris c.t. Eleocharis palustris c.t. Wm05 Typha latifolia c.t. Wm06 Scirpus acutus c.t. Wm07 Juncus balticus c.t. Wm50 Carex sitchensis c.t. Carex sitchensis c.t. Wm51 Dulichium arundinaceum c.t. Ws06 Salix sitchensis c.t. Ws50 Spiraea douglasii c.t./Sphagnum; Spiraea c.t. Ws52 Alnus rubra – Lysichiton americanum c.t. Ws53 Thuja plicata – Tsuga heterophylla/Lysichiton americanum c.t.

* Community type

252 Wetlands of British Columbia: A Field Guide to Identiﬁcation  . Wetland Site Associations corresponding with classifications in Washington and Montana

Lincoln County,Washington (Crawford 2000) Montana (Hansen et al. 1995) Fl01 Alnus incana c.t. Fl02 Mountain alder/red-osier dogwood assoc. Fl03 Salix lasiandra c.t. Fl05 Salix drummondiana/Calamagrostis canadensis h.t.** Fl06 Sandbar willow assoc. Salix exigua c.t. Fl07 Water birch/Wood’s rose c.t.* Fm01 Black cottonwood/Common snowberry forest Gs01 Saltgrass – Clustered field sedge c.t. Distichilis spicata h.t. Gs02 Hordeum jubatum c.t. Gs04 Tufted hairgrass assoc. Deschampsia cespitosa h.t. Wb02 Salix planifolia/Carex aquatilis h.t. Wf01 Carex rostrata h.t. and Carex aquatilis h.t. Wf02 Betula nana/Carex rostrata h.t. Wf05 Carex lasiocarpa h.t. Wf08 Carex limosa h.t. Wf09 Eleocharis pauciflora h.t. Wm01 Northwest Territory sedge assoc. Wm02 Equisetum fluviatile h.t. Wm04 Common spikerush assoc. Eleocharis palustris h.t. Wm05 Typha latifolia h.t. Wm06 Hard-stem bulrush assoc. Scirpus acutus h.t. Wm07 Baltic rush – Silverweed cinquefoil Juncus balticus c.t. c.t. Ws03 Salix bebbiana c.t. Ws04 Salix drummondiana/Carex rostrata h.t.

* Community type ** Habitat type

Appendix 4 253 APPENDIX 5 Scientific names with common names of species used in this guide

Abies amabilis amabilis fir Abies lasiocarpa subalpine fir Acer glabrum Douglas maple Acer macrophyllum bigleaf maple Achillea millefolium yarrow Actaea rubra baneberry Agrostis aequivalvis Alaska bentgrass Agrostis exarata spike bentgrass Agrostis stolonifera creeping bentgrass Alnus viridis ssp. crispa green alder Alnus incana mountain alder Alnus rubra red alder Alopecurus aequalis little meadow-foxtail Amphiscirpus nevadensis Nevada bulrush Andromeda polifolia bog-rosemary Angelica lucida seacoast angelica Antennaria pulcherrima showy pussytoes Arctostaphylos uva-ursi kinnikinnick Aster ciliolatus Lindley’s aster Aster conspicuus showy aster Aster ericoides tufted white prairie aster Aster subspicatus Douglas’ aster Athyrium filix-femina lady fern Atriplex patula common orache Aulacomnium palustre glow moss Barbilophozia sp. leafy liverwort Betula nana scrub birch Betula occidentalis water birch Betula papyrifera paper birch Blechnum spicant deer fern Bolboschoenus maritimus seacoast bulrush Brachythecium spp. feathermosses Brasenia schreberi water shield Bryum pseudotriquetrum marsh thread-moss Calamagrostis canadensis bluejoint Calamagrostis stricta slimstem reedgrass Calliergon sp. water-moss Calliergon stramineum straw spear-moss Caltha leptosepala white mountain marsh-marigold Campylium sp. star-moss Campylium stellatum yellow star-moss Campylopus atrovirens bristly swan-neck moss

254 Wetlands of British Columbia: A Field Guide to Identiﬁcation Carex anthoxanthea yellow-flowered sedge Carex aquatilis water sedge Carex atherodes awned sedge Carex chordorrhiza cordroot sedge Carex disperma soft-leaved sedge Carex exsiccata inflated sedge Carex lanuginosa woolly sedge Carex lasiocarpa slender sedge Carex leptalea bristle-stalked sedge Carex limosa shore sedge Carex livida pale sedge Carex lyngbyei Lyngbye’s sedge Carex magellanica poor sedge Carex obnupta slough sedge Carex pauciflora few-flowered sedge Carex pluriflora many-flowered sedge Carex praegracilis field sedge Carex sitchensis Sitka sedge Carex tenuiflora sparse-leaved sedge Carex utriculata beaked sedge Ceratophyllum sp. hornwort Ceratophyllum demersum common hornwort Chamaecyparis nootkatensis yellow-cedar Chamaedaphne calyculata leatherleaf Chara sp. stonewort Cicuta douglasii Douglas’ water-hemlock Circaea alpina enchanter’s-nightshade Cladina spp. reindeer lichens Cladonia spp. clad lichens Comarum palustre marsh cinquefoil Conioselinum gmelinii Pacific hemlock-parsley Coptis trifolia three-leaved goldthread Cornus canadensis bunchberry Cornus stolonifera red-osier dogwood Deschampsia cespitosa ssp. tufted hairgrass beringensis Deschampsia cespitosa ssp. tufted hairgrass cespitosa Deschampsia cespitosa tufted hairgrass Distichlis spicata seashore saltgrass Distichlis spicata var. spicata seashore saltgrass Distichlis spicata var. stricta alkali saltgrass Drepanocladus sp. hook-mosses Drepanocladus aduncus common hook-moss Drosera anglica great sundew Drosera rotundifolia round-leaved sundew

Appendix 5 255 Dulichium arundinaceum three-way sedge Eleocharis palustris common spike-rush Eleocharis quinqueflora few-flowered spike-rush Elliottia pyroliflorus copperbush Elodea canadensis Canadian waterweed Elymus glaucus blue wildrye Elymus trachycaulus slender wheatgrass Empetrum nigrum crowberry Epilobium angustifolium fireweed Equisetum sp. horsetail Equisetum arvense common horsetail Equisetum fluviatile swamp horsetail Equisetum hyemale scouring-rush Equisetum pratense meadow horsetail Equisetum telmateia giant horsetail Erigeron peregrinus subalpine daisy Eriophorum angustifolium narrow-leaved cotton-grass Eriophorum chamissonis Chamisso’s cotton-grass Fauria crista-galli deer-cabbage Festuca rubra red fescue Fragaria virginiana wild strawberry Fritillaria camschatcensis northern rice-root Fucus sp. brown seaweed Galium trifidum small bedstraw Gaultheria hispidula creeping-snowberry Gaultheria shallon salal Geum macrophyllum large-leaved avens Glaux maritima sea-milkwort Glyceria borealis northern mannagrass Glyceria elata tall mannagrass Gymnocarpium dryopteris oak fern Heracleum maximum cow-parsnip Homatocaulis vernicosus stick hook-moss Hordeum brachyantherum meadow barley Hordeum jubatum foxtail barley Hylocomium splendens step moss Hypericum anagalloides bog St. John’s-wort Iris pseudacorus yellow iris Isoetes echinospora bristle-like quillwort Juncus arcticus arctic rush Juncus balticus Baltic rush Juniperus communis common juniper Kalmia microphylla western bog-laurel Eurhynchium praelongum slender beaked-moss Kobresia myosuroides Bellard’s kobresia Koeleria macrantha junegrass

256 Wetlands of British Columbia: A Field Guide to Identiﬁcation Larix laricina tamarack Lathyrus palustris marsh peavine Ledum groenlandicum Labrador tea Lemna sp. duckweed Leptarrhena pyrolifolia leatherleaf saxifrage Leymus mollis dune wildrye Ligusticum scoticum beach lovage Lilaeopsis occidentalis western lilaeopsis Lobelia dortmanna water lobelia Lonicera involucrata black twinberry Lupinus nootkatensis Nootka lupine Lysichiton americanus skunk cabbage Lythrum salicaria purple loosestrife Maianthemum dilatatum false lily-of-the-valley Maianthemum stellatum star-flowered false solomon’s-seal Malus fusca Pacific crab apple Matteuccia struthiopteris ostrich fern Meesia triquetra three-ranked hump-moss Menyanthes trifoliata buckbean Menziesia ferruginea false azalea Mnium spp. leafy mosses Muhlenbergia richardsonis mat muhly Myrica gale sweet gale Myriophyllum spicatum Eurasian water-milfoil Myriophyllum sp. water-milfoil Myriophyllum verticillatum verticillate water-milfoil Najas flexilis wavy water nymph Nuphar lutea yellow pond-lily Nuphar lutea ssp. polysepala yellow pond-lily Oemleria cerasiformis Indian-plum Oenanthe sarmentosa Pacific water-parsley Oplopanax horridus devil’s club Osmorhiza berteroi mountain sweet-cicely Oxycoccus oxycoccos bog cranberry Phalaris arundinacea reed canarygrass Philonotis fontana spring moss Phragmites australis common reed Picea X spruce Picea mariana black spruce Picea sitchensis Sitka spruce Pinus contorta lodgepole pine Pinus contorta var. contorta shore pine Pinus contorta var. latifolia lodgepole pine Pinus monticola western white pine Plantago macrocarpa Alaska plantain Plantago maritima sea plantain

Appendix 5 257 Platanthera dilatata fragrant white rein orchid Pleurozium schreberi red-stemmed feathermoss Poa pratensis Kentucky bluegrass Poa secunda ssp. juncifolia Nevada bluegrass Poa secunda Sandberg’s bluegrass Poa trivialis rough bluegrass Polygonum amphibium water smartweed Polystichum munitum sword fern Populus balsamifera balsam poplar Populus balsamifera ssp. black cottonwood trichocarpa Populus tremuloides trembling aspen Potamogeton sp. pondweed Potamogeton amplifolius large-leaved pondweed Potamogeton foliosus closed-leaved pondweed Potamogeton gramineus grass-leaved pondweed Potamogeton natans floating-leaved pondweed Potamogeton praelongus long-stalked pondweed Potamogeton pusillus small pondweed Potamogeton richardsonii Richardson’s pondweed Potamogeton robbinsii Robbin’s pondweed Potentilla anserina common silverweed Potentilla egedii coast silverweed Potentilla gracilis graceful cinquefoil Pteridium aquilinum bracken fern Ptilium crista-castrensis ostrich-plume feather-moss Puccinellia nuttalliana Nuttall’s alkaligrass Puccinellia pumila dwarf alkaligrass Pyrola asarifolia pink wintergreen Racomitrium lanuginosum hoary rock-moss Ranunculus aquatilis white water-buttercup Ranunculus flammula lesser spearwort Ranunculus orthorhynchus straight-beaked buttercup Rhynchospora alba white beak-rush Rhytidiadelphus loreus lanky moss Rhytidiadelphus squarrosus bent-leaf moss Ribes bracteosum stink currant Rosa acicularis prickly rose Rosa nutkana Nootka rose Rosa sp. rose Rosa woodsii prairie rose Rubus chamaemorus cloudberry Rubus parviflorus thimbleberry Rubus spectabilis salmonberry Ruppia maritima widgeon-grass Salicornia virginica American glasswort

258 Wetlands of British Columbia: A Field Guide to Identiﬁcation Salix barclayi Barclay’s willow Salix bebbiana Bebb’s willow Salix brachycarpa short-fruited willow Salix candida sage willow Salix commutata under-green willow Salix drummondiana Drummond’s willow Salix exigua sandbar willow Salix glauca grey-leaved willow Salix lucida ssp. lasiandra Pacific willow Salix lucida Pacific willow Salix maccalliana MacCalla’s willow Salix myrtillifolia bilberry willow Salix pedicellaris bog willow Salix planifolia tea-leaved willow Salix prolixa Mackenzie willow Salix sitchensis sitka willow Sambucus racemosa red elderberry Sanguisorba canadensis sitka burnet Sanguisorba officinalis great burnet Sarracenia purpurea common pitcher-plant Scheuchzeria palustris scheuchzeria Schoenoplectus acutus hard-stemmed great bulrush Schoenoplectus americanus American bulrush Schoenoplectus pungens sharp bulrush Schoenoplectus subterminalis water clubrush Schoenoplectus tabernaemontani soft-stemmed great bulrush Scirpus microcarpus small-flowered bulrush Scorpidium sp. sausage-moss Scorpidium revolvens rusty hook-moss Scorpidium scorpioides hooked scorpion-moss Senecio triangularis arrow-leaved groundsel Siphula ceratites northern waterfingers Sium suave hemlock water-parsnip Sparganium angustifolium narrow-leaved bur-reed Sparganium natans small bur-reed Spartina gracilis alkali cordgrass Spergularia canadensis Canadian sand-spurry Sphagnum sp. peat-moss Sphagnum angustifolium poor-fen peat-moss Sphagnum austinii tough peat-moss Sphagnum capillifolium common red peat-moss Sphagnum fuscum common brown peat-moss Sphaghum magellanicum Magellanic peat-moss Sphagnum papillosum fat peat-moss Sphagnum Group I peat-mosses, Group I Sphagnum Group II peat-moss - Group II

Appendix 5 259 Sphagnum Group III peat-mosses, Group III Sphagnum Group IV peat-mosses, Group IV Spiraea douglasii pink spirea Stachys mexicana Mexican hedge-nettle Streptopus amplexifolius clasping twistedstalk Streptopus lanceolatus rosy twistedstalk Stuckenia pectinata fennel-leaved pondweed Suaeda calceoliformis seablite Subularia aquatica awlwort Symphoricarpos albus common snowberry Taraxacum officinale common dandelion Thalictrum occidentale western meadowrue Thuja plicata western redcedar Tiarella trifoliata foamflower Triantha glutinosa sticky false-asphodel Tomentypnum sp. fen moss Tomentypnum nitens golden fuzzy fen moss Triantha glutinosa sticky false-asphodel Trichophorum alpinum Hudson bay clubrush Trichophorum cespitosum tufted clubrush Trifolium wormskioldii springbank clover Triglochin maritima seaside arrow-grass Trollius albiflorus globeflower Tsuga heterophylla western hemlock Tsuga mertensiana mountain hemlock Typha latifolia common cattail Urtica dioica stinging nettle Utricularia gibba humped bladderwort Utricularia intermedia flat-leaved bladderwort Utricularia macrorhiza greater bladderwort Utricularia sp. bladderwort Vaccinium sp. blueberry Vaccinium alaskaense/ Alaska/oval-leaved blueberry ovalifolium Vaccinium uliginosum bog blueberry Vaccinium vitis-idaea lingonberry Vahlodea atropurpurea mountain hairgrass Valeriana sitchensis Sitka valerian Veratrum viride Indian hellebore Veronica scutellata marsh speedwell Viburnum edule highbush-cranberry Viola palustris marsh violet Warnstorfia sp. hook-mosses Zostera marina common eel-grass

260 Wetlands of British Columbia: A Field Guide to Identiﬁcation APPENDIX 6 Wildlife species Affiliated with Wetlands and Related Ecosystems

The following list includes wildlife species that commonly live in or use wetlands and related ecosystems. Amphibians Mole Salamanders Ambysomatidae Northwestern Salamander Ambystoma gracile Long-toed Salamander Ambystoma macrodactylum Tiger Salamander Ambystoma tigrinum Giant Salamanders Dicamptodontidae Paciﬁc Giant Salamander Dicamptodon tenebrosus Lungless Salamanders Plethodontidae Wandering Salamander Aneides vagrans Ensatina Ensatina eschscholtzii Coeur d’Alene Salamander Plethodon idahoensis Western Red-backed Salamander Plethodon vehiculum Newts Salamandridae Rough-skinned Newt Taricha granulosa Tailed Frogs Ascaphidae Coastal Tailed Frog Ascaphus truei Rocky Mountain Tailed Frog Ascaphus montanus Spadefoots Pelobatidae Great Basin Spadefoot Spea intermontana True Toads Bufonidae Western Toad Bufo boreas Treefrogs Hylidae Paciﬁc Treefrog Hyla regilla Boreal Chorus Frog Pseudacris maculata True Frogs Ranidae Red-legged Frog Rana aurora Bullfrog Rana catesbeiana Green Frog Rana clamitans Columbia Spotted Frog Rana luteiventris Northern Leopard Frog Rana pipiens Oregon Spotted Frog Rana pretiosa Wood Frog Rana sylvatica

Appendix 5 261 Reptiles Western Pond Turtle Clemmys marmorata Painted Turtle Chrysemys picta Rattlesnake Crotalus viridis Gopher Snake Pituophis melanoleucus Common Garter Snake Thamnophis sirtalis Western Terrestrial Garter Snake Thamnophis elegans Birds Loons Gaviidae Red-throated Loon Gavia stellata Paciﬁc Loon Gavia paciﬁca Common Loon Gavia immer Grebes Podicipedidae Pied-billed Grebe Podilymbus podiceps Horned Grebe Podiceps auritus Red-necked Grebe Podiceps grisegena Eared Grebe Podiceps nigricollis Western Grebe Aechmophorus occidentalis Clark’s Grebe Aechmophorus clarkii Pelicans Pelecanidae American White Pelican Pelecanus erythrorhynchos Bitterns and Herons Ardeiadae American Bittern Botaurus lentiginosus Great Blue Heron Ardea herodias Great Egret Casmerodius albus Cattle Egret Bubulcus ibis Green Heron Butorides striatus Black-crowned Night Heron Nycticorax nycticorax Swans, Geese, and Ducks Anatidae Tundra Swan Cygnus columbianus Trumpeter Swan Cygnus buccinator Mute Swan Cygnus olor Greater White-fronted Goose Anser albifrons Snow Goose Chen caerulescens Brant Branta bernicla Canada Goose Branta canadensis Wood Duck Aix sponsa Green-winged Teal Anas crecca

262 Wetlands of British Columbia: A Field Guide to Identification American Black Duck Anas rubripes Mallard Anas platyrhynchos Northern Pintail Anas acuta Blue-winged Teal Anas discors Cinnamon Teal Anas cyanoptera Northern Shoveler Anas clypeata Gadwall Anas strepera Eurasian Wigeon Anas penelope American Wigeon Anas americana Canvasback Aythya valisineria Redhead Aythya americana Ring-necked Duck Aythya collaris Greater Scaup Aythya marila Lesser Scaup Aythya affinis Northen Long-tailed Duck Clangula hyemalis Common Goldeneye Bucephala clangula Barrow’s Goldeneye Bucephala islandica Bufflehead Bucephala albeola Hooded Merganser Lophodytes cucullatus Common Merganser Mergus merganser Red-breasted Merganser Mergus serrator Ruddy Duck Oxyura jamaicensis Rails and Coots Rallidae Virginia Rail Rallus limicola Sora Porzana carolina American Coot Fulica americana Cranes Gruidae Sandhill Crane Grus canadensis Hawks and Eagles Accipitridae Osprey Pandion haliaetus Bald Eagle Haliaeetus leucocephalus Grouse and Ptarmigans Phasianidae Blue Grouse Dendragapus obscurus Willow Ptarmigan Lagopus lagopus Ruffed Grouse Bonasa umbellus Sharp-tailed Grouse Tympanuchus phasianellus ssp. columbianus

Appendix 6 263 Plovers Charadriidae Semipalmated Plover Charadrius semipalmatus Killdeer Charadrius vociferus Lesser Golden-Plover Pluvialis dominica Stilts and Avocets Recurvirostridae American Avocet Recurvirostra americana Sandpipers Scolopacidae Greater Yellowlegs Tringa melanoleuca Lesser Yellowlegs Tringa ﬂavipes Solitary Sandpiper Tringa solitaria Willet Catoptrophorus semipalmatus Spotted Sandpiper Actitis macularia Upland Sandpiper Bartramia longicauda Hudsonian Godwit Limosa haemastica Semipalmated Sandpiper Calidris pusilla Western Sandpiper Calidris mauri Least Sandpiper Calidris minutilla Baird’s Sandpiper Calidris bairdii Pectoral Sandpiper Calidris melanotos Dunlin Calidris alpina Sanderling Calidris alba Stilt Sandpiper Calidris himantopus Buff-breasted Sandpiper Tryngites subruﬁcollis Short-billed Dowitcher Limnodromus griseus Long-billed Dowitcher Limnodromus scolopaceus Common Snipe Gallinago gallinago Wilson’s Phalarope Phalaropus tricolor Red-necked Phalarope Phalaropus lobatus Gulls and Terns Laridae Bonaparte’s Gull Larus philadelphia Mew Gull Larus canus Ring-billed Gull Larus delawarensis California Gull Larus californicus Herring Gull Larus argentatus Franklin’s Gull Larus pipixcan Forster’s Tern Sterna forsteri Black Tern Chlidonias niger Caspian Tern Sterna caspia Common Tern Sterna hirundo

264 Wetlands of British Columbia: A Field Guide to Identification Owls Strigidae Great Gray Owl Strix nebulosa Great Horned Owl Bubo virginianus Northern Hawk Owl Surnia ulula Kingfishers Alcedinidae Belted Kingfisher Ceryle alcyon Tyrant Flycatchers Tyrannidae Alder Flycatcher Empidonax alnorum Willow Flycatcher Empidonax trailii Eastern Kingbird Tyrannus tyrannus Waxwings Bombycillini Bohemian Waxwing Bombycilla garrulus Cedar Waxwing Bombycilla cedrorum Dippers Cinclidae American Dipper Cinclus mexicanus Wrens Troglodytidae House Wren Troglodytes aedon Winter Wren Troglodytes troglodytes Marsh Wren Cistothorus palustris Chickadees Paridae Black-capped Chickadee Parus atricapillus Swallows Hirundinidae Purple Martin Progne subis Tree Swallow Tachycineta bicolor Violet-green Swallow Tachycineta thalassina Northern Rough-winged Swallow Stelgidopteryx serripennis Bank Swallow Riparia riparia Cliff Swallow Hirundo pyrrhonota Barn Swallow Hirundo rustica Kinglets and Thrushes Muscicapidae Ruby-crowned Kinglet Regulus calendula Veery Catharus fuscescens Gray-cheeked Thrush Catharus minimus Hermit Thrush Catharus guttatus

Appendix 6 265 Warblers Sylviidae American Redstart Setophaga ruticilla Ovenbird Seiurus aurocapillus Northern Waterthrush Seiurus noveboracensis Common Yellowthroat Geothlypis trichas Yellow-rumped Warbler Dendroica coronata Yellow Warbler Dendroica petechia Palm Warbler Dendroica palmarum MacGillivray’s Warbler Oporornis tolmiei Wilson’s Warbler Wilsonia pusilla Yellow-breasted Chat Icteria virens Sparrows and Blackbirds Fringillinae American Tree Sparrow Spizella arborea Chipping Sparrow Spizella passerina Savannah Sparrow Passerculus sandwichensis Fox Sparrow Passerella iliaca Song Sparrow Melospiza melodia Lincoln’s Sparrow Melospiza lincolnii Sharp-tailed Sparrow Ammodramus caudacutus Swamp Sparrow Melospiza georgiana Yellow-headed Blackbird Xanthocephalus xanthocephalus Red-winged Blackbird Agelaius phoeniceus Bobolink Dolichonyx oryzivorus Rusty Blackbird Euphagus carolinus Brewer’s Blackbird Euphagus cyanocephalus Mammals Shrews Soricidae Black-backed Shrew Sorex arcticus Paciﬁc Water Shrew Sorex bendirii Pygmy Shrew Sorex hoyi Dusky Shrew Sorex monticolus Water Shrew Sorex palustris Bats Chiroptera Big Brown Bat Eptesicus fuscus Spotted Bat Euderma maculatum Silver-haired Bat Lasionycteris noctivagans Western Red Bat Lasiurus blossevillii ssp. frantzi Hoary Bat Lasiurus cinereus

266 Wetlands of British Columbia: A Field Guide to Identiﬁcation California Myotis Myotis californicus Western Small-footed Myotis Myotis ciliobrum Western Long-eared Myotis Myotis evotis Keen’s Long-eared Myotis Myotis keenii Little Brown Myotis Myotis lucifugus Northern Long-eared Myotis Myotis septentrionalis Yuma Myotis Myotis yumanensis Townsend’s Big-eared Bat Plecotus townsendii Mountain Beavers Aplodontiidae Mountain Beaver Aplodontia rufa Beavers Castoridae Beaver Castor canadensis Voles and Lemmings Arvicolidae Southern Red-backed Vole Clethrionomys gapperi Northern Red-backed Vole Clethrionomys rutilus Water Vole Microtis richardsonii Meadow Vole Microtis pennsylvannicus Townsend’s Vole Microtis townsendii Long-tailed Vole Microtis longicaudus Brown Lemming Lemmus sibiricus Northern Bog Lemming Synaptomys borealis Muskrat Ondatra zibethicus Jumping Mice Zapodidae Western Jumping Mouse Zapus princeps Meadow Jumping Mouse Zapus hudsonius Weasels Mustelidae Wolverine Gulo gulo Mink Mustela vison River Otter Lontra canadensis Striped Skunk Mephitis mephitis Ermine Mustela erminea Long-tailed Weasel Mustela frenata Racoon Procyonidae Raccoon Procyon lotor

Appendix 6 267 Bears Ursidae Black Bear Ursus americanus Grizzly Bear Ursus arctos Deer Cervidae Moose Alces alces Elk Cervus elaphus Mule Deer Odocoileus hemionus Caribou Rangifer tarandus

268 Wetlands of British Columbia: A Field Guide to Identiﬁcation APPENDIX 7 Glossary

Aerobic Occurring in the presence of free oxygen, either as a gas in the atmosphere or dissolved in water.

Alkaline Water or soil with a pH greater than 7.4. Relatively high concentration of available base cations.

Anaerobic Occurring in conditions devoid of oxygen.

Annual ﬂood Flooding occurs at least once per year.

Biogeoclimatic subzone A climatic region characterized by a distinct climax plant association on zonal sites.

Biogeoclimatic zone A climatic region with similar broad macroclimate characterized by a distinct zonal plant order.

Brown mosses A guild of peatland mosses usually indicating mineral rich site conditions. Includes Campylium stellatum, Drepanocladus spp., Scorpidium spp., and Tomenthypnum nitens.

Canopy Cover of branches and leaves formed collectively by the crowns of trees, shrubs, or other plants.

Capillary In a soil, the ﬁne spaces between soil particles.

Capillary action Particles attract soil moisture, and surface tension is strong enough to cause moisture to rise up through the soil, above the watertable.

Class, site Ecosystems with broadly similar vegetation physiognomy (or species guild), hydrology, and water quality ( 1997).

Clayey Predominant textural classes are clay, silty clay, sandy clay, or clay loam.

Climax community A self-perpetuating community whose species composition is expected to be relatively stable and long lasting.

Closed basin or pond Basin receives water from surrounding upland only, no inlet or outlet channel.

Appendix 6 269 Coarse water sedges Large, broad-leaved sedge species including Carex aquatilis, C. atherodes, C. exsiccata, C. sitchensis, and C. utriculata.

Common Occurs frequently, and representative ecosystems are readily found, but it is not a predominant association of the region (Steen and Roberts 1988).

Common species Species that can occur in a Site Association, but do not deﬁne the community. They usually have a presence >30% and a cover >1%.

Constant species Species that occur in a classification unit with relatively high frequency but low mean cover that may help to define the community. They are defined as having presence of >66% and cover <10%.

Diagnostic species A species that occurs primarily within a single classiﬁ- cation unit.

Diatomaceous earth Composed mainly of the siliceous shells of diatoms. It is frequently more nearly mineral than organic in composition.

Disclimax A self-perpetuating community that strongly differs in species composition from the edaphic or climatic climax expected for the site; normal succession has been arrested by an external physical or anthropogenic factor. Results from changes to physical characteristics of the site, associated with disturbances such as ﬁre, intensive grazing, or avalanche (Province of British Columbia 1998).

Dominant species The strucurally most dominant species within a site or the species that contributes greatest vegetation cover to the community.

Drawdown Decrease in water level of lakes or steams, exposing substrate that is normally submerged.

Dwarf shrubs Plants with woody stems that are generally less than 15 cm tall at maturity. Andromeda polifolia, Arctostaphylos uva-ursi, Em- petrum nigrum, Gaultheria hispidula, Kalmia microphylla, Linnaea borealis, Oxycoccus oxycoccos, Rubus chamaemorus, Rubus pedatus, Vaccinium caespitosum, and Vaccinium vitis-idaea are the most common wetland dwarf shrub species.

270 Wetlands of British Columbia: A Field Guide to Identification Emergents Upright plants rooted in water or exposed to seasonal flooding, emerging above water surface. Does not include some submergents that normally lie entirely under water but have flowering parts that break the surface. Includes mostly sedges, rushes, bulrushes, and other grass-like forbs.

Ericaceous shrubs Shrubs of family Ericaceae. Andromeda, Chamaedaphne, Gaultheria, Kalmia, Ledum, Oxycoccus, and Vaccinium are the most common wetland genera.

Eutrophic Very rich nutritional status, abundant supply of nutrients.

Feathermosses Upland moss species with a feather-like form including Hylocomium splendens, Pleurozium schreberi, and Ptilium crista- castrensis.

Fibric Poorly decomposed peat with large amounts of well-preserved ﬁbre readily identiﬁable as to botanical origin.

Flark Elongated wet depressions separated by raised ribs in patterned peatlands. The long axis is always perpendicular to the direction of waterﬂow.

Floating mat Mat of peat held together by roots and rhizomes underlain by water or fluid, loose peat ( 1988). Floating-leaved plants Rooted or free-floating plants with leaves normally floating on water surface.

Flooding Surface inundation by moderate to fast moving water. Usually associated with sedimentation and erosion (see also Inundation).

Fluvial Sites occurring along ﬂowing watercourses, the watercourse itself, and the surrounding (riparian) terrain and vegetation. Subject to ﬂooding and sedimentation processes (Province of British Colum- bia 1998).

Forb Any non-graminoid herb species.

Forested Sites with >10% canopy cover of tree species >10 m tall (see also Treed).

Frequent ﬂooding Flood return interval of 2–5 years.

Appendix 7 271 Gleyed A soil condition resulting from prolonged soil saturation, which is manifested by the presence of bluish or greenish colours throughout the soil mass or in mottles (usually orange spots or streaks).

Graminoid Plants with a grass-like growth form including rushes (Jun- caceae), grasses (Poaceae), and sedges (Cyperaceae).

Groundwater Water passing through or standing in soil and underlying strata. Free to move by gravity ( 1988).

Herb Non-woody vascular plants. Includes forbs and graminoids.

Hollow 1. A wet depression or pool found between hummocks or mounds. 2. A sunken basin or depression, often sloped and having an outﬂow. Includes gullies with slow streams where there is little sedimentation or erosion.

Humic Highly decomposed organic material. Small amounts of ﬁbre can be identiﬁed to botanical origin ( 1988).

Hummock A mound composed of organic material, often composed of Sphagnum peat (see also Mound). Slight hummocks are 0.3–1 m tall and spaced >7 m apart. Moderate hummocks are 0.3–1 m tall and spaced 3–7 m apart. Strong hummocks are 0.3–1 m tall and spaced 1–3 m apart.

Humus Dead and decaying organic material at the soil surface.

Hydric 1. A site where water is removed so slowly that the watertable is at or above the soil surface all year. 2. A Gleysol or Organic soil.

Hydrogeomorphic classiﬁcation Classiﬁcation of wetland and riparian ecosystems based on hydrological and geomorphological features and processes.

Hydrophytic plant species Any plant adapted for growing on permanently saturated soils deﬁcient in oxygen.

Hygric Water removed slowly enough to keep soil wet for most of the growing season; permanent seepage and mottling usually below 30 cm in depth.

Hypereutrophic Sites with very high salinity or alkalinity.

272 Wetlands of British Columbia: A Field Guide to Identiﬁcation Inundation Surface ﬂooding by standing or slow-moving water.

Lacustrine Sites adjacent to lakes and ponds directly affected by lake wave action, sedimentation, and ﬂooding.

Lagg Depressed margin of a bog or fen; generally wetter than surrounding area, often contains open water.

Lawn Relatively ﬂat expanse of wetland moss usually raised above water level. Contrast with Hummock and Hollow.

Lifeform A plant growth form that displays an obvious relationship to important environmental factors (Mueller-Dombois and Ellen- berg 1974).

Limnic material Composed of coprogenous earth (sedimentary peat), diatomaceous earth, or marl.

Linked basin Basin receives water from upland and an inflow stream; excess water flows through an outflow. Includes basins with slow streams where there is little sedimentation or erosion (Province of British Columbia 1998).

Loamy Textural classes are loam and sandy loam (Steen and Roberts 1988).

Marl Sediments composed of shells of aquatic animals and CaCO3 pre- cipitated in water.

Mesic 1. Organic material in an intermediate stage of decomposition where some fibres can be identified as to botanical origin. 2. Medi- um soil moisture regime where a site has neither excess soil moisture nor a moisture deficit.

Microtopography Small-scale (i.e., < 2 m) variations in soil surface elevation (e.g., hummocks and hollows).

Minerotrophic Nourished by mineral water. Refers to wetlands that receive nutrients from ﬂowing or percolating mineral groundwater ( 1988).

Minerotrophic indicator species Plant species requiring relatively high concentrations of nutrients associated with mineral groundwater.

Appendix 7 273 Mire British term embracing all kinds of peatlands and peatland vegetation (modiﬁed from  1988).

Moderately acidic Having a soil pH value of 4.5–5.5.

Moist No water deﬁcit occurs. Current need for water does not exceed supply; temporary groundwater table may be present (Pojar et al. 1987).

Montane A high-elevation region occurring below the subalpine.

Mound Mounds composed of mineral materials (see also Hummock).

Muskeg Algonquin term for peatland. Usually applied to areas with Sphagnum mosses, tussocky sedges, and an open growth of scrub- by trees (modiﬁed from  1988).

Neotenous An animal that is sexually mature in the larval stage.

Neutral pH Having a soil pH value between 6.5 and 7.4. Available base cation concentration is high enough to buffer acidic conditions.

Occasional ﬂooding Flood interval greater than 5 years.

Oligotrophic Relatively poor in nutrients.

Ombrotrophic Nourished by rain. Peatlands entirely dependent on nutrients deposited by precipitation ( 1988).

Overﬂow basin Basin receives water from upland only; excess water ﬂows through an outlet channel (Province of British Columbia 1998).

Paludiﬁcation Succession or conversion of upland or mineral wetland habitats to peatland through accumulation of peat.

Palustrine Basins, depressions, slopes, and small waterbodies with a continually high watertable and poor-drainage wetland landscape units.

Palustrine hollow Hollow receives groundwater from upslope; excess water ﬂows through channel or watertrack.

Patterned peatland Peatlands marked by distinct patterns of vegetation in alternating raised ridges and depressions (ﬂark) forms. Sites are slightly sloping and ridges form perpendicular to the direction of waterﬂow.

274 Wetlands of British Columbia: A Field Guide to Identiﬁcation Peat Partly decomposed plant material deposited under saturated soil conditions.

Peatland A generic term including all types of peat-covered terrain. Many peatlands are a complex of swamps, bogs, and fens, sometimes called a “mire complex” ( 1988).

Physiognomic Referring to vegetation structure or strata.

Rarely ﬂooded Flooding occurs only during extreme events.

Riparian Along the bank of a river, lake, or wetland.

Saline The presence of soluble salts in the soil parent material at concentrations that affect plant growth.

Sandy Textural classes are loamy sand and sand (Steen and Roberts 1988).

Saturated A soil condition in which all voids (pore spaces) between soil particles are ﬁlled with water.

Sedimentary peat (coprogenous earth) Peat formed beneath a body of standing water composed of aquatic plant debris modiﬁed by aquatic animals. Material is loosely consolidated, slightly sticky, dark brown to black, and usually well decomposed (humic). Syn- onyms: aquatic peat, loonshit, allochthonous peat, detrital peat, gyttja ( 1988).

Seepage Groundwater discharge having less ﬂow than a spring.

Shrub Perennial plants usually with more than one low-branching woody stem and <10 m tall.

Silty Predominant textural classes are silt and silt loam (Steen and Roberts 1988).

Site Association A group of related ecosystems physically and biologically similar enough that they have or would have similar vegetation at climax (Meidinger and Pojar 1991).

Slightly acidic Having a soil pH value of 5.5–6.5.

Stand A plant community that is relatively uniform in composition, structure, and habitat conditions.

Appendix 7 275 Subassociation Subunits of an association that are floristically very similar but distinguished by the predominance of one to several differentiating species.

Subhydric Soil moisture regime where water is removed slowly enough to keep watertable at or near the surface for most of the year; permanent seepage 0–30 cm below surface.

Subhygric Water removed slowly enough to keep soil wet for a signiﬁ- cant part of the growing season; some temporary seepage and possibly mottling below 20 cm.

Submergents Plants that normally lie entirely beneath water. Some species have ﬂowering parts that break the water surface.

Succession Replacement of one community by another; often progresses to a stable terminal community called the climax.

Treed Sites with >10% canopy cover of tree species >2 m and <10 m tall (see also Forested).

Tussock A thick tuft of sedge or other vegetation forming a small mound of solid ground in a wetland ( 1988).

Uncommon Occurs infrequently in a region (Steen and Roberts 1988).

Very acidic Having a soil pH value less than 4.5. Low concentration of available base cations.

Very moist Rooting-zone groundwater present during the growing season (water supply exceeds demand). Groundwater table >30 cm deep (Pojar et al 1987).

Very wet Groundwater table at or above the ground surface throughout most of the growing season. von Post A qualitative scale of peat decomposition.

Watertable The upper surface of the zone of saturation within the soil proﬁle.

Wet Rooting-zone groundwater present during the growing season (water supply exceeds demand). Groundwater table >0 cm but < 30 cm below soil surface (Pojar et al 1987).

276 Wetlands of British Columbia: A Field Guide to Identiﬁcation Wetland Sites dominated by hydrophytic vegetation where soils are water-saturated for a sufﬁcient length of time such that excess water and resulting low soil oxygen levels are principal determinants of vegetation and soil development.

Wetland complex Consists of two or more wetland communities occurring in close proximity in the same system and inﬂuenced or linked by the same moisture and nutrient regime.

Appendix 7 277 LITERATURE CITED

Agriculture Canada Expert Committee on Soil Survey. 1987. The Canadi- an system of soil classification. Supply and Services Canada, Ottawa, Ont. Agric. Can. Publ. 1646. Anderson, L. E. 1990. A checklist of Sphagnum in North America north of Mexico. The Bryologist 93(4): 500–501. Anderson, L. E., H. A. Crum, and W. R. Buck. 1990. List of mosses of North America north of Mexico. The Bryologist 93(4): 448–499. Annas, R. M. 1974. Boreal ecosystems of the Fort Nelson area of northeastern British Columbia. M.Sc. thesis. Univ. British Columbia, Vancouver, B.C. Banner, A. 1983. Classification and successional relationships of some bog and forest ecosystems near Prince Rupert, British Columbia. M.Sc. thesis. Univ. British Columbia, Vancouver, B.C. Banner, A., W. MacKenzie, S. Haeussler, S. Thomson, J. Pojar, and R. Trowbridge. 1993. A field guide to site identification and interpretation for the Prince Rupert Forest Region. B.C. Min. For., Res. Br., Victoria, B.C. Land Manage. Handb. 26. Banner, A., J. Pojar, and R. Trowbridge. 1986. Representative wetland types of the northern part of the Pacific Oceanic Wetland Region. B.C. Min. For., Res. Br., Victoria, B.C. Beil, C. E. 1969. The plant associations of the Cariboo – Aspen – Lodge- pole Pine – Douglas-fir parkland zone. Ph.D. dissertation. Univ. British Columbia, Vancouver, B.C. Belland, R. J. and D.H. Vitt. A key to British Columbia wetland bryophytes based mostly on field characteristics. Unpublished. Boe, J. S. 1994. Nest site selection by Eared Grebes in Minnesota. Condor 96: 19–35. Bridgham, S. D., J. Pastor, J. A. Janssens, C. Chapin, and T. J. Malterer. 1996. Multiple limiting gradients in peatlands: a call for a new par- adigm. Wetlands 16(1): 45–65.

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280 Wetlands of British Columbia: A Field Guide to Identification Delong, D., A. McLeod, A. MacKinnon, and L. Jang. 1990. A field guide for identification and interpretation of ecosystems of the north- east portion of the Prince George Forest Region. B. C. Min. For., Victoria, B.C. Land Manage. Handb. 22. Douglas, G. W., D. Meidinger, and J. Pojar. 1998. Illustrated flora of British Columbia: gymnosperms and dicotyledons (Aceraceae through Asteraceae). Volume 1. B.C. Min. Sustainable Resource Management and B.C. Min. For., Victoria, B.C. ———. 1999. Illustrated flora of British Columbia: dicotyledons (Dia- pensiaceae through Onagraceae). Volume 3. B.C. Min. Sustainable Resource Management and B.C. Min. For., Victoria, B.C. ———. 1999. Illustrated flora of British Columbia: dicotyledons (Ora- banchaceae through Rubiaceae). Volume 4. B.C. Min. Sustainable Resource Management and B.C. Min. For., Victoria, B.C. ———. 2000. Illustrated flora of British Columbia: dicotyledons (Sali- caceae through Zygophyllaceae) and Pteridiophytes. Volume 5. B.C. Min. Sustainable Resource Management and B.C. Min. For., Victoria, B.C. ———. 2001. Illustrated flora of British Columbia: monocotyledons (Acoraceae through Najadaceae). Volume 6. B.C. Min. Sustainable Resource Management and B.C. Min. For., Victoria, B.C. ———. 2001. Illustrated flora of British Columbia: monocotyledons (Orchidaceae through Zosteraceae). Volume 7. B.C. Min. Sustain- able Resource Management and B.C. Min. For., Victoria, B.C. ———. 2002. Illustrated flora of British Columbia: general summary, maps, and keys. Volume 8. B.C. Min. Sustainable Resource Man- agement and B.C. Min. For., Victoria, B.C. Douglas, G. W., G. B. Straley, D. Meidinger, and J. Pojar. 1998. Illustrat- ed flora of British Columbia: gymnosperms and dicotyledons (Balsaminaceae through Cuscutaceae). Volume 2. B.C. Min. Sus- tainable Resource Management and B.C. Min. For. Victoria, B.C. Earnst, S. L. 1994. Tundra Swan habitat preferences during migration in North Dakota. J. Wildl. Manage. 58(3): 546–551.

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282 Wetlands of British Columbia: A Field Guide to Identification Harris, A. G., S. C. McMurray, P. W. C. Uhlig, J. K. Jeglum, R. F. Foster, and G. D. Racey. 1996. Fieldguide to the wetland ecosystem classification for Northwestern Ontario. Ont. Min. Nat. Resour., Northwest Sci. Tech., Thunder Bay, Ont.  Field Guide FG-01. Harrison, C. 1978. A field guide to the nests, eggs and nestlings of North American birds. Collins, Toronto, Ont. Hartman, G. F. and T. G. Brown. 1988. Forestry-fisheries planning considerations on coastal floodplains. The Forestry Chronicle. Howes, D., M. Morris, and M. Zacharias. 1999. British Columbia estuary mapping system Version 1. Resources Inventory Committee, Vic- toria, B.C. Johnson, D., L. Kershaw, A. MacKinnon, and J. Pojar. 1995. Plants of the western boreal forest and aspen parkland. Lone Pine Publishing, Edmonton, Alta. Johnston, C. A. and R. J. Naiman. 1990. Aquatic patch creation in relation to beaver population trends. Ecology 71(4): 1617–1621. Kauffman, J. B. 1982. Synecological effects of cattle grazing riparian ecosystems. M.Sc. thesis. Oregon State Univ., Corvallis, Oreg. Kauffman, J. B. and W. C. Krueger. 1984. Livestock impacts on riparian ecosystems and streamside management implications...a review. J. Range Manage. 37(5): 430–438. Klinka, K., H. Qian, and A. Ceska. Provisional classification of non- forested plant communities in coastal British Columbia. Unpublished. Luttmerding, H. A., A. D. Demarchi, E. C. Lea, D. V. Meidinger, and T. Vold. 1990. Describing ecosystems in the field. 2nd ed. B.C. Min. Environ., Lands, and Parks and B.C. Min. For., Victoria, B.C.  Manual 11. MacKenzie, W. H. and R. Klassen. 1999. User guide for 97 ver. 2.0. Web document .

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284 Wetlands of British Columbia: A Field Guide to Identification Mueller-Dombois, D. and H. Ellenberg. 1974. Aims and methods of vegetation ecology. Wiley, New York, N.Y. National Wetlands Working Group (). 1988. Wetlands of Cana- da. Environ. Can., Sustainable Development Br., Ottawa, Ont. Ecological Land Classification Series 24. Newman, D. G. and C. R. Griffen. 1994. Wetland use by River Otters in Massachusetts. J. Wildl. Manage. 58(1): 18–23. Noss, R. F. 1994. At what scale should we manage biodiversity? College of Forestry, Univ. Idaho, Moscow, Idaho. Päivänen, J. 1997. Forested mires as a renewable resource—toward a sustainable forestry practice. In Northern forested wetlands: ecology and management. C. C. Trettin, M. F. Jurgensen, D. F Grigal, M. R. Gale, and J. K. Jeglum (editors). Lewis Publishers, New York, N.Y., pp. 27–44. Peden, D. G. 1967. Vegetation and ecology of Rithet’s Bog, Royal Oak, British Columbia. BSc. Honours thesis. Univ. Victoria, Victoria, B.C. Peterson, E. B., N. M. Peterson, and D. S. McLennan. 1996. Black cottonwood and balsam poplar managers’ handbook for British Columbia. For. Can. and B.C. Min. For., Victoria, B.C.  Rep. 250. Platts, W. S. 1982. Sheep and cattle grazing strategies on riparian-stream environments. In Wildlife-Livestock relationships symposium: Proc.10. Univ. Idaho, Forest, Wildlife, and Range Experimental Station, Moscow, Idaho. Pojar, J., K. Klinka, and D. V. Meidinger. 1987. Biogeoclimatic ecosystem classification in British Columbia. For. Ecol. and Manage. 22: 119–154. Powell, G. W., K. J. Cameron and R. F. Newman. 2000. Analysis of livestock use of riparian areas: literature review and research needs assessment for British Columbia. B.C. Min.For. Res. Br., Victoria, B.C. Work. Pap 52.

Literature cited 285 Province of British Columbia.1998. Field manual for describing terrestrial ecosystems. B.C. Min. Environ., Lands and Parks and B.C. Min.For., Victoria, B.C. Land Manage. Handb. 25. Reid, F. A. 1993. Managing wetlands for waterbirds. Trans. North Ameri- ca Wildlife and Natural Resource Conf. 58: 345–350. Resource Inventory Committee (). 1998. Inventory methods for pond-breeding amphibians and Painted Turtle. B.C. Min. Envi- ron., Lands, and Parks, Victoria, B.C. ———. 1999. Inventory methods of waterfowl and allied species: Loons, Grebes, Swans, Geese, Ducks, American Coot and Sandhill Crane. Ver. 2.0. B.C. Min. Environ., Lands and Parks, Victoria, B.C. Revel, R. D. 1972. Phytogeocoenoses of the Sub-Boreal Spruce biogeoclimatic zone in north central British Columbia. Ph.D. dissertation. Univ. British Columbia, Vancouver, B.C. Richards, L. A. 1969. Diagnosis and improvement of saline and alkali soils. , Soil and Water Conserv. Res. Br., Washington, D.C. Richter, K. O. and A. L. Azous. 1995. Amphibian occurrence and wetland characteristics in the Puget Sound Basin. Wetlands 15(3): 305–312. Roberts, A. 1984. Guide to wetland ecosystems of the Sub-Boreal Spruce A subzone. Cariboo Forest Region, British Columbia. B.C. Min. For., Res. Section, Williams Lake, B.C. ———. 1986. Willows of the lower Skeena and Bulkley valleys and their relative importance as winter food for Moose. B.C. Min. Environ., Wildl. Br., Smithers, B.C. Runka, G. G. and T. Lewis. 1981. Preliminary wetland managers manual for the Cariboo Resource Management Region. B.C. Min. Envi- ron., Terrestrial Studies Br., Victoria, B.C. Semlitsch, R. D. and R. Bodie. 1998. Are small, isolated wetlands expend- able? Conserv. Biol. 12(5): 1129–1133. Sims, R. A. and K. A. Baldwin. Sphagnum species in northwestern On- tario: a ﬁeld guide to their identiﬁcation. Can. For. Serv., Sault Ste. Marie, Ont. / Tech. Rep. TR-30.

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Literature cited 287 near Houston, the patterning seems to reflect wave refraction after waves pass through a constriction. Elevated ribs form perpendicular to gradual water flow in patterned fens. In this dramatic example from

A northern fen in fall colours (Stewart-Cassiar highway at the Yukon border)

Wetlands and related ecosystems are important components of British Columbia's ecological diversity. This guide describes over 100 distinct wetland, riparian floodplain, estuarine, shallow-water, and "transition" site associations. Each association is described in a one-page fact-sheet summary that outlines its essential environmental and biological attributes. B.C. Government Publication Services, Publisher www.publications.gov.bc.ca/ Please visit our website for information on acquiring copies of this Land Management Handbook, and other publications published by the Government of British Columbia.

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